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Annals of Tropical Medicine 
and Parasitology 




I 


I 



THE UNIVERSITY OF LIVERPOOL 


Annals 

OF 

Tropical Medicine and 
Parasitology 

ISSUED BY THE 

Liverpool School of Tropical Medicine 


Edited by 

Professor J. W. W. STEPHENS, M.D.Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 


VOLUME XIII 

(May 12, 1919, to March 15, 1920) 

fVith Frontispiece, twelve plates, forty-six figures in text, thirty-one charts, 

and three maps 


LIVERPOOL: 

AT THE UNIVERSITY PRESS, 57 ASHTON STREET 



L U ( r 


I'd.',/ i; 

'VI 


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PUBLIC HFA1.;■* 


X 



CONTENTS 


No. i. May 12, 1919 

PAGE 

Smith, A. Malins. 

A Contribution to the Question of the Number of Races in the species Entamoeba 

histolytica . 1 

Matthews, J. R. 

The Course and Duration of an Infection with Entamoeba colt ... ... ... 17 

Macfie, J. W. S. 

Two Parasites of Naja nigricollis . 23 


Evans, Alwen M. 

On the Genital Armature of the Female Tsetse-flies ( Glossina ) . 31 

Yorke, Warrington ; and Macfie, J. W. S. 

Strongylidae in Horses : VII.— Cylicostomum pateratum sp. n. 57 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.; Macfie, J. W. S.; 

Cooper, C. Forster ; and Carter, H. F. 

Studies in the Treatment of Malaria : XXII.—Intramuscular Injections 
of Quinine Bihydrochloride Grains 15 on each of two consecutive 
dap only, in Malignant Tertian Malaria . 63 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.*; Macfie, J. W. S.; 

Cooper, C. Forster ; and Carter, H. F. 

Studies in the Treatment of Malaria: XXIII.—Oral Administration of 
Quinine Sulphate Grains*30 on each of two consecutive days weekly, 
over a period of five weeks, in Malignant Tertian Malaria . 69 


Stephens, Lieut.-Col. J. W. W. ; Yorke, VV. ; Blacklock, B.; Macfie, J. W. S,; 
Cooper, C. Forster ; and Carter, H. F. 

Studies in the Treatment of Malaria : XXIV.—The Disappearance of 

Crescents under Quinine Treatment. 73 

Stephens, Lieut.-Col. J. W. W. ; Yorke, W. ; Blacklock, B.; Macfie, J. W. S.; 
Cooper, C. Forster ; and Carter, H. F. 

Studies in the Treatment of Malaria : XXV.—Arsenic in Malignant Tertian 

Malaria . 75 


Matthews, J. R.; and Smith, A. Malins. 

The Intestinal Protozoal Infections among Convalescent Dysenteries examined 

at the Liverpool School of Tropical Medicine. (Third Report) ... ... 83 

Matthews, J. R.; and Smith, A. Malins. 

The Spread and Incidence of Intestinal Protozoal Infections in the Population 
of Great Britain : IV.—Asylum Patients. V.—University and School 

Cadets. 91 


v 



CONTENTS 
No. 2. July 3i, 1919 

Macfie, Dr. J. W. S. page 

Presentation of the Mary Kingsley Medal . 95 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.; and Macfie, J. W. S. 

Studies in the Treatment of Malaria : XXVI.—The Action of Arsenic and of 

Quinine on Quartan Malaria. 97 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.; and Macfie, J. VV. S. 

Studies in the Treatment of Malaria : XXVII.—Intravenous Injections of 
Novarsenobillon and Intramuscular Injections of Quinine Bihydrochloride 
in Simple Tertian Malaria . 101 

Rodhain, J. 

Filaria pertenue , n.sp., Provoquant une Dermofilariose Cheloi'diforme chez 

Cephalophus sylvicultor 109 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.; Macfie, J. W. S.; 
and O’Farrell, Capt. W. R. 

Studies in the Treatment of Malaria : XXV III.—Quitenine Hydrochloride 

in Simple Tertian Malaria . 117 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B. ; Macfie, J. W. S.; 
and O’Farrell, Capt. W. R. 

Studies in the Treatment of Malaria : XXIX.—Oral Administration of 
Liquor Arsenicalis minims 30 daily for 16 days with Quinine Bihydro¬ 
chloride grains* 15 Intramuscularly on the first and second, eighth and 
ninth, fifteenth and sixteenth days, in Simple Tertian Malaria ... 119 

Stephens, Lieut.-Col. J. W. W.; Yorke, W.; Blacklock, B.; Macfie, J. W. S.; 
and O’Farrell, Capt. W. R. 

Studies in the Treatment of Malaria : XXX.—At what time after Cessation 
of Quinine Treatment do Relapses occur in Simple Tertian Malaria ? 


(Second communication). 125 

Yorke, W.; and Macfie, J. W. S. 

The Phagocytosis of Erythrocytes by an Amoeba of the Limax Type. 133 

Yorke, W.; and Macfie, J. W. S. 

Strongylidae in Horses : VIII.—Species found in American Horses . 137 

Lipkin, I. J. 

On the Distribution and Destruction of Quinine in Animal Tissues . 149 

Smith, A. Malins. 


Cases of Acute Amoebic Dysentery in Asylum Patients never out of England... 177 
Blacklock, B. ; and Carter, H. F. 

The Experimental Infection, in England, of Anopheles plumbeus , Hal., with 

Plasmodium vivax (Sporozoites in Salivary Glands). (Preliminary Note) 187 

Blacklock, B. ; and O’Farrell, Capt. W. R. 

Note on a Case of Multiple Infection by Dr a run cuius medinensis ... ... 189 


vi 



CONTENTS 


No. 3. December 10, 1919 

PAG£ 

Scon’, Henry Harold. 

Coincident Malaria and Enteric Fever . 195 

Young, William John. 

The Metabolism of White Races living in the Tropics: II.—The composition 

of the Urine . 215 

Hatori, Juro. 

On the Endemic Tsutsugamushi Disease of Formosa . ... 233 

Fielding, J. W. 

Notes on the Bionomics of Stegomyia fas data y Fabr. (Parti) ... ... 259 

Blacklock, B. 

Ancylostoma ceylaniaim in the Cat in Durban . 297 


vii 



CONTENTS 


No. 4. March 15, 1920 


Carter, H. R. 

The Mechanism of the Spontaneous Elimination of Yellow Fever from Endemic 
Centres. 

Young, W. J. 

The Metabolism of White Races living in the Tropics: 111 .—"Flic Influence' 
of External Temperature and Rate of Cooling upon the Respiratory 
Metabolism . 

Morris, Hubert M. 

The Hypopus of Carpoglyphus attonymus, Haller 

Macfie, J. W. S. 

Xerophthalmia in a Native of the Gold Coast 

Macfie, J. W. S. 

An Observation on the Effect of Malaria in Leukaemia. 

Breinl, A.; and Young, W. J. 

Tropical Australia and its Settlement . 

Blacklock, B.; and Carter, Henry F. 

The Experimental Infection in England of Anopheles plumbeus , Stephens, and 
Anopheles bifurcatus 9 L., with Plasmodium vivax . 

Blacklock, B.; and Carter, Henry F. 

Observations on Anopheles ( Coelodiazesis) plumbeus 9 Stephens, with special 
reference to its Breeding-places, Occurrence in the Liverpool District, 
and Possible Connection with the Spread of Malaria . 

Carter, Henry F. 

Descriptions of the Male Genital Armatures of the British Anopheline 
Mosquitoes . . 

Yorke, Warrington. 

On Human Trypanosomiasis in Peru . 


viii 


PAGE 

299 

313 

339 

343 

347 

3 S 1 

4*3 

421 

453 

459 






Volume XIII 


May, 1919 


No. 1 


ANNALS 

OF 

TROPICAL MEDICINE AND 
PARASITOLOGY 


ISSUED BY 

THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE 


Edited by 

«» 

Professor J. W. W. STEPHENS, M.D. Cantab., D.P.H. 

Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S. 
Professor WARRINGTON YORKE, M.D. 



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Professors - - JOHN WILLIAM WATSON STEPHENS, M.D., 

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The following courses of instruction will be given by the 
Liverpool School of Tropical Medicine during 1919 •— 

Full Course begins 6 January. Advanced Course begins 1 June. 
Diploma Examination, 2 April. Certificate Examination, 1 July. 

Full Course begins 16 September. 

Diploma Examination, 9 December. 

These dates are subject to revision. 

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For prospectus and further information, application should be made 
to the Dean of the Medical Faculty, University of Liverpool. 


The following have obtained the Diploma in Tropical Medicine of 
the University of Liverpool: — 


Diploma in Tropical Medicine 


Data of 
Diploma 

1904 Augustine, Henry Joshua 
1904 Bennett, Arthur King 
1904 Bruce, William James 
1904 Byrne, John Scott 
1904 Clayton, Thomas Morrison 
1904 Dalziel, John McEwen 
1904 Dee, Peter 
1904 Greenidge, Oliver Campbell 
1904 Hehir, Patrick 
1904 Khan, Saiducsafor 
1904 Laurie, Robert 
1904 Madurkin, Alfred Robert 
1904 McConnell, Robert Ernest 
1904 Nicholson, James Edward 
1904 Philipson, Nicholas 
1904 Sharraan, Eric Harding 
1904 Thomson, Frank Wyville 

1904 Walker, George Francis Clegg 

1905 Anderson, Catherine EtxnsHe 
1905 Brown, Alexander 

1905 Caldwell, Thomas Cathcart 
1905 Critien, Attifco 
1905 Hooton, Alfred 
1905 Hudson, Chaika Tllson 
1905 Illingtoa, Edmund Moritz 


Date of 
Diploma 

1905 Macfariane, Robert Maxwell 
1905 Maddock, Edward Cecil Gordon 
1905 Moore, James Jackson 
1905 Nightingale, Samuel Shore 
1905 Radcliffe, Percy Alexander Hunt 

1905 Young, John Cameron 

1906 Adic, Joseph Rosamond 
1906 Arnold, Frank Arthur 
1906 Bate, John Brabant 
1906 Bennetts, Harold Graves 
1906 Carter, Robert Markham 
1906 Chisholm, James Alexander 
1906 Clements, Robert WilHam 
1906 Dundas, James 

1906 Faichnie, Norman 
1906 Jeffreys, Herbert Castelman 
1906 Mackenzie, Donald Francis 
1906 Pailthorpe, Mary Elizabeth 
1906 Palmer, Haroid Thombury 
1906 Pearse, Albert 
1906 Sampey, Alexander William 
1906 Smithson, Arthur Ernest 
1906 Taylor, Joseph van Soraeron 
1906 Taylor, WilHam Irwin 
1906 Tynan, Edward Joseph 



Date of 
Diploma 

1906 Watson, Cecil Francis 
1906 Willcocks, Roger Durant 

1906 Williamson, George Alexander 

1907 Allan, Alexander Smith 
1907 Allwood, James Aldrcd 
1907 Bond, Ashton 

1907 Branch, Stanley 
1907 Collinson, Walter Julius 
1907 Davey, John Bernard 
1907 Donaldson, Anson Scott 
1907 Fell, Matthew Henry Gregson 
1907 Gann, Thomas William Francis 
1907 Graham, James Drummond 
1907 Hiscock, Robert Carroll 
1907 Keane, Joseph Gerald 
1907 Kennan, Richard Henry 
1907 Kcnrick, William Hamilton 
1907 Le Fanu, George Ernest Hugh 
1907 Mackey, Charles 
1907 Maddox, Ralph Henry 
1907 McCarthy, John McDonald 
1907 Raikes, Cuthbert Taunton 
1907 Ryan, Joseph Charles 
1907 Vallance, Hugh 


1908 Caverliill, Austin Mack 

1908 Crawford, Gilbert Stewart 

1908 Dalai, Kaikhusroo Rustomji 

1908 Dansey-Browning, George 

1908 Davidson, James 

1908 Dickson, John Rhodes 

1908 Dowdall, Arthur Melville 

1908 Glover, Henry Joseph 

1908 Greaves, Francis Wood 

1908 Goodbody, Cecil Maurice 

1908 Harrison, James Herbert Hugh 

1908 Joshi, Lemuel Lucas 

1908 Le Fanu, Cecil Vivian 

1908 Luethgen, Carl Wilhelm Ludwig 

1908 Mama, Jamshed Byramji 

1908 McCay, Frederick William 

1908 McLellan, Samuel Wilson 

1908 Pearce, Charles Ross 

1908 Schoorel, Alexander Frcdcrik 

1908 Smith, John Macgregor 

1908 Stewart, George Edward 

1908 Tate, Gerald William 

1908 Whyte, Robert 


1909 Abercrombie, Rudolph George 
1909 Allin, John Richard Percy 
1909 Armstrong, Edward Randolph 
1909 Barrow, Harold Percy Waller 
1909 Beatty, Guy 
1909 Carr-White, Percy 
1909 Chevallier, Claude Lionel 
1909 Clark, William Scott 
1909 Cope, Ricardo 
1909 Fleming, William 
1909 Hanschcll, Hother McCormick 
1909 Hayward, William Davey 
1909 Henry, Sydney Alexander 
1909 Innes, Francis Alexander 


Date of 
Diploma 

1909 Jackson, Arthur Frame 
1909 Kaka, Sorabji Manekji 
1909 McCabe-Dallas, Alfred Alexander 
Donald 

1909 Meldrum, William Percy 
1909 Murphy, John Cullman 
1909 Samuel, Mysore Gnananandaraju 
1909 Shroff, Kawasjee Byramjee 
1909 Thornely, Michael Harris 
1909 Turkhud, Violet Ackroyd 
1909 Webb, William Spinks 
1909 Yen, Fu-Chun 


1910 Brabaxon, Edward 

1910 Castellino, Louis 

1910 Caul crick, James Akilade 

1910 Dowden, Richard 

1910 Haigh, William Edwin 

1910 Hamilton, Henry Fleming 

1910 Hefferman, William St. Michael 

1910 Hipwell, Abraham 

1910 Homer t Jonathan 

19101 Houston, William Mitchell 

1910 James, William Robert Wallace 

1910 Johnstone, David Patrick 

1910 Korke, Vishnu Tatyaji 

1910 Macdonald, Angus Graham 

1910 Macfie, John Wm. Scott 

1910 Manuk, Mack Walter 

1910 Murison, Cecil Charles 

1910 Nanavati, Kishavlal Balabhai 

1910 Nauss, Ralph Welty 

1910 Oakley .Philip Douglas 

1910 Pratt, Ishmael Charles 

1910 Sabastian, Thiruchelvam 

1910 Shaw, Hugh Thomas 

1910 Sieger, Edward Louis 

1910 Sousa, Pascal John de 

1910 Souza, Antonio Bernardo de 

1910 Waterhouse, John Howard 

1910 White, Maurice Forbes 


1911 Blacklock, Breadalbane 
1911 Brown, Frederick Forrest 
1911 Chand, Diwan Jai 
1911 Holmes, John Morgan 
1911 levers, Charles Langley 
1911 lies, Charles Cochrane 
1911 Ingram, Alexander 
1911 Kirkwood, Thomas 
1911 Knowles, Benjamin 
1911 Liddle, George Marcus Berkeley 
1911 Lomas, Emanuel Kenworthy 
1911 Mackarell, William Wright 
1911 MacKnight, Dundas Simpson 
1911 Mascarenhas, Joseph Victor 
1911 Murray, Ronald Roderick 
19U Oluwole, Akidiya Ladapo 
1911 Rao, Koka Ahobala 
1911 Sinton, John Alexander 
1911 Tarapurvalla, Byramji Shavakshah 
1911 Taylor, John Archibald 
1911 Woods, William Medlicott 



Data of 
Diploma 

1912 Aeria, Joseph Reginald 

1912 Anderson, Edmund Litchfield 

1912 Borle, James 

1912 Bowie, John Tait 

1912 Brassey, Laurence Percival 

1912 Christie, David 

1912 Dillon, Henry de Courcy 

1912 Dunn, Lillie Eleanor 

1912 Hardwicke, Charles 

19x2 Jagose, Jam shed Rustomji 

1912 Kochhar, Mela Ram 

1912 McGusty, Victor William Tighe 

1912 Milne, Arthur James 

1912 Mitra, Manmatha Nath 

1912 Myles, Charles Duncan 

1912 Pelly, Huntly Nevins 

1912 Prasad, Bindeshwari 

1912 Prentice, George 

1912 Ross, Frank 

1912 Russell, Alexander James Hutchison 

1912 Ruthven, Morton Wood 

1912 Sandilands, John 

1912 Seddon, Harold 

1912 Smalley, James 

1912 Strickland, Percy Charles Hutchison 

1912 Watson, William Russel 


X913 Austin, Charles Miller 

1913 Banker, Shiavux Sorabji 

1913 Becker, Johann Gerhard us 

1913 Carrasco, Milton 

1913 Clark, James McKillican 

1913 Forsyth, Charles 

1913 Grahame, Malcolm Claude Russell 

1913 Grieve, Kelburnc King 

1913 Hargreaves, Alfred Ridley 

19x3 Hepper, Evelyn Charles 

1913 Hiranand, Pandit 

1913 Jackson, Oswald Egbert 

1913 Khaw, Ignatius Oo Kek 

1913 MacKelvie, Maxwell 

1913 MacKinnon, John MacPhail 

1913 Macmillan, Robert James Alan 

1913 Mouat-Biggs, Charles Edward Forbes 

1913 Noronha, John Carmel 

1913 O’Connor, Edward 

1913 Olubomi-Beckley, Emanuel 

1913 Pcstonji, Ardcshir Behramshah 

1913 Puttanna, Dodballapur Sivappa 

1913 Reford, John Hope 

1913 Smith, Edward Arthur 

1913 Stewart, Samuel Dudley 

1913 Walker, Frederick Deardcn 


Da to of 
Diploma 

1913 Wilbe, Ernest Edward 
19x3 Wilson, Hubert Francis 
1913 Yin, Ulg Ba 
1913 Young, William Alexander 


1914 Arculli, Hassan el 
1914 Chohan, Noormahomed Kasembha 
1914 Connell, Harry Bertram 
i.914 Gerrard, Herbert Shaw 
1914 Gimi, Hirji Dorabji 
1914 Gwynnc, Joseph Robert 
1914 Hodkinson, Samuel Paterson 
1914 Jackson, Arthur Ivan 
1914 Kaushash, Ram Chander 
1914 Kelsall, Charles 
1914 Luanco y Cuenca, Maximino 
1914 Misbah, Abdul-Ghani Naguib 
1914 Naidu, Bangalore Pasupulati 
Balakrishna 

1914 Rowe, John Joseph Stephen 

1914 Roy, Raghu Nath 

1914 Shiveshwarkar, Ramchandra Vishnu 

1914 Sur, Sachindra Nath 

1914 Talati, Dadabhai Cursedji 

1914 Wilkinson, Arthur Geden 

1914 Wright, Ernest Jenner 

1915 Lobo, John Francis 
1915 Madhok, Gopal Dass 
1915 Pearson, George Howorth 
1915 Swami, Karumuri Virabhadra 

1915 Wood, John 

1916 Barseghian, Mesroob 
1916 Chaliha, Lakshmi Prasad 
1916 Lim, Albert Liat Juay 
1916 Lim, Harold Liat Hin 
1916 Metzger, George Nathaniel 
1916 Soderstrom, Erik Daniel 

1916 Wheeler, Louis 

19x7 Chapman, Herbert Owen 

1917 Krishnamoorthy, Yedatore Venkoba 
19x7 Lipkin, Isaac Jacob 

1918 Watts, Rattan Claud 

1919 Celestin, Louis Abel 

1919 Cummings, Eustace Henry Taylor 
1919 Darling, Georgina Renington 
1919 Maplestone, Philip Alan 
1919 Rustomjee, Khusshuyee Jamesidjce 
1919 Turner, Gladys Maude 



ANNALS OF TROPICAL MEDICINE 
AND PARASITOLOGY 


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Robinson (1914) says the reverse.’ The references should be 
collected in alphabetical order of authors’ surnames at the end of 
the paper, and arranged in the following way: — 

Robinson, S. (1914). 1 The spleen in malaria.* Annals of Nosology , 

Vol. XX, pp. 20-25. 

Smith, T. (1900). ‘ Enlargement of the spleen in malaria.* Journal of 

Patnomttry y Vol. I, pp. 1-20. 

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A CONTRIBUTION TO THE QUESTION 
OF THE NUMBER OF RACES IN THE 
SPECIES ENTAMOEBA HISTOLYTICA 

BY 

A. MALINS SMITH, M.A., Cantab. 

From the Liverpool School of Tropical Medicine 
{Received for publication January 31, 1919) 

In the course of a recent paper (Smith, 1918) I discussed 
tentatively the idea put forward by Wenyon and O’Connor (1917) 
and by Dobell and Jepps (1918) that the species Entamoeba 
histolytica is composed of races distinguishable from each other by 
the dimensions of their cysts. I gave reasons for the conclusion that 
two races, the 'small* (average diameter 77 m) and the 'ordinary* 
(average diameter 12*6/1) certainly exist, but that the evidence for 
further races was insufficient. It was shown that the proof of the 
existence of such further races depended upon the measurement of 
large samples of a considerable % number of infections. In my 
measurements the samples taken were too small, being usually fewer 
than fifty cysts, whereas Dobell and Jepps (1918) measured very 
large samples (five hundred cysts), but the number of infections they 
examined (seven) was small. 

In order tb obtain more complete data for the solution of this 
question I have measured fairly large samples (one hundred at least) 
of a considerable number of infections, twenty in all, and the results 
are set forth in the present paper. In addition to these samples of 
100 or more, samples of fifty to one hundred were measured in 
thirteen cases, and some evidence is also drawn from these 
measurements. All the measurements are of cysts belonging to the 
'ordinary* strain, no 'small* cysts having been measured. The 
methods of measurement were the same as those used in my previous 
paper (Smith, 1918). 

Before the results of these measurements can be accurately 
considered, a preliminary question must be dealt with, namely 



whether the average size of the cysts of an infection remains constant 
from one day to another. Dobell and Jepps (1918) say ‘For the 
complete demonstration of this fact (the existence of strains), 
however, it is necessary to prove that the mean diameter of the cysts 
from any patient is not subject to any considerable variation from 
day to day, but remains constant.' They state that this constancy is 
a fact which is confirmed daily in the course of routine examinations. 
Measurements in support of it are brought forward only in one case, 
their E42, in which five hundred cysts measured on May 21st, 1917, 
gave, a mean diameter of 1299/1, five hundred cysts measured on 
June 23rd, 1917, had a mean diameter of 13*24/1, giving a difference 
of 0*25/1. Some evidence will be brought forward later as to whether 
this is really a negligible difference in so large a sample. At present 
it may be remarked that the evidence by actual measurement is‘only 
given for one infection, and can scarcely therefore sufficiently establish 
the theory of general constancy in the average size of the cysts of 
infections. It is necessary to measure the cysts of a larger number 
of infections for this purpose. In Table I are shown the results 
of the measurement of samples of one hundred cysts (with two 
exceptions, viz., seventy-five and seventy-eight respectively) on 
different dates in eight different infections. 

It will be seen that while the agreement between the results on 
different dates is in some cases close, in others it is by no means 
so, and it is not possible to tell by mere inspection what conclusion 
should be drawn from the results. It is first of all necessary 
to know the error of sampling, i.e. what variations are to be 
expected in measurements of random samples of one hundred 
from the same stool. The following data refer chiefly to the 
results of measurements of successive samples of fifty cysts from 
the same stool, but it will be seen later that deductions 
can be drawn from them as to samples of one hundred cysts. 
In every case where one hundred cysts were measured the difference 
between the average size of the first fifty and the second 
fifty was recorded. In cases where two hundred cysts were 
measured from the same stool there were, of course, four lots of fifty, 
and this gave six different numbers for the differences between 
fifties. Where three hundred were examined containing six fifties, 
there were fifteen differences to be obtained. In this way from 



3 


Tablk I. 




1 Dates of 


i 


Case 

Date of stool | 

emetine 

No. 

I Average diam. (//) 

Differences (fi) 


: 

treatment 

measured 

1 of each 100 cysts 

Q 

27.7.18 

1.6.18 

300 

1 

1 12.83. 12*56, 1253 

c*33, o*o6, 0*03 



to 


1 



15.8.18 

10.10.18 

12.6.18 

(inclusive) 

100 

! 

1 

0*38, o*i 1, 0*08 



LOO 

12-50 

0*05 

II 

3.5.18 

3.5.18 

200 

12*24, 12*o6 

0*29, 0*11, 0*10 



to 


I 

0*08, 0* 19 


14.5.18 

14.5.18 

IOO 

"•95 

119,1*08 



(inclusive) 


1*00, 0*90 


21.5.18 


IOO 

j "'14 

1*40, 1*59 

1*30 , 107 


30.7.18 


3 00 

i 3 * 3 »> 1 3 * 54 ? 13*4 

1*48 , 1*17 

1*36, 1*25 

16 

19.4.18 

19.4.18 

100 

1 3**3 

0*02 



to 





7.5.18 

30.4.18 
(inclusive) 

IOO 

* 3‘*5 


IO 

3.4.18 

7.4.18 

IOO 

1312 

0*00 



to 





19.4.18 

18.4.18 

(inclusive) 

100 

13*12 

0*85 


30*4*18 


IOO 

12*27 

0*76 

1 

! 

4.5.18 


IOO 

12*36 

0*09 

*7 1 

4.4.18 

j 

25.3.18 1 

* 100 

I 3 ‘ 3 I 

o*o 7 



to 





9.4.18 

5.4.18 

(inclusive) 

IOO 

12*91 

0*40 


25.4.18 


IOO 

13*24 

0*33 

T 3 

5.4.18 

25.3.18 

IOO 

12*80 




to 





6.4.18 

5.4.18 

(inclusive) 

IOO 

1 

* 3-27 

0*47 

12 

4.3.18 

23.1.18 1 

75 

12*60 

0*57 



to 

1 




30.4.18 

1.2.18 

100 

* 3**7 


1 


(inclusive) 

1 

1 J 

1 


*5 

26.3.18 

25.3.18 

IOO 


C*II 



to 1 





27.3.18 

5.4.18 

(inclusive) 

7 8 1 

« 3>4 




4 


eighteen infections sixty-eight different numbers were obtained 
showing the differences found between the average sizes of random 
samples of fifty cysts from the same stool. The figures are set out 
in Table II, and are placed in order of magnitude in Table III. 


Table II. 


Case 

Date of stool 

Differences between 5o’s 
from same stool ( ju ) 

Differences between loo’s 
from same stool (/ i ) 

i 

22.5.18 

°'34 


3 

23.10.18 

0*03 


4 

22.7.18 

0*10 


5 

15.4.1? 

0-02 


6 

7.9.18 

0*20 


7 

7 - 5 - 1 8 

0-07 


S 

29.8.18 

0-24 


9 

27.7.18 

15.8.18 
IO.lO.l8 

0*00, 0-02, 0-05, 0*05, 0*05, 
0*10, 0*10, 0*10, 0*12, 
0-12, 0-17, 0*20, 0-20, 
0*22, 0*22 

0-03 

0-37 

o*o2, 0-15. 0*17 

lo 

3.4.18 

19.4.18 

30.4.18 

4.5.18 

0-05 

0*03 

015 

0*41 


ii 

3.5.18 

14.5.18 

21.5.18 

30.7.18 

0*03, 0-08, 0-12, 0*20, 0-24. 
0-32 

0*10 

0-24 

0*01, 0*02, 0-15, 0-17, 0*17, 
0*18, 0-19, 0*20, 0-27, 
0-28, 0-34, 0-35, 0 - 45 , 
0*47, 0-62 

o*i8 

0-40, 0-17, 0-23 

12 

30.4.18 

0*10 


*3 

5.4.18 

6.4.18 

0*22 

0-34 


*4 

28.1I.l8 

0-41 


*5 

26.3.18 

0-17 


16 

I9.4.18 

7.5.18 

0*17 

0-05 


l 7 

4.4.18 

9.4.18 
25.4.18 

0*03 

0-20 

0*41 


iS 

II.12.l8 

0*00, 0-03, 0*27, 0-27, 0-31, 
0-31 

0*03 

*9 

j 16.5.18 

0-32 



5 


Table III. 



Differences between 50*8 in ft 
in order of magnitude 

Differences between ioo’s in fi 
in order of magnitude 

o*io and below 

0*00, 0*00, 0*01, 0*02, 0*02, 0'02, 

0-03, 0*03, 0-03, 0*03, 0*03, 0-03, 
0*05, 0*05, 0*05, 0-05, 0-05, 0-07, 
0'08, 0*10, o-lo, O’lO, O'IO, O' 10, 
O'lO 

0-02, 0-03 

O' I I to 0*20 

1 

0*12, 0-12, 0*12, 0-15, 0-15, 0*17, 

0*17, 0*17, 0*17, 0*17, 0*18, 0-19, 

0-15, 0-17, 0*17, o-i8 


1 0*20, 0-20, 0-20, Q'20, 0*20, 0*20 


0*21 to 0-30 j 

i 

j 

0*22, 0*22, 0*22, 0'24, 0*24, 0*24, 

0*27, 0*27, 0-27, 0*28 

023 

0*31 to 0-40 j 

1 

0-31, 0-31, 0-32, 0-32, oj4, 0-34, 

°’ 34 > o -3 5 . °-37 

0-40 

0*41 to 0*50 | 

1 

0-41, 0*41, 0-41, o*45, 0-47 

— 

0-51 to o*6o | 

! 

— 

— 

0*6! to 070 1 

! 

0-62 

— 


From this it may be seen that only one of the sixty-eight differences 
exceeds 0*60/i, only six of the sixty-eight exceed 0*40/1, and half 
are below 018 /1. Taking ten chances out of eleven as high 
probability, there is high probability that the differences between two 
random samples of fifty will not exceed 0*40/1. Now for samples of 
one hundred cysts this probability will be greater still. Table III 
shows also the differences between random samples of one hundred. 
Of these there are eight, and it will be seen that not one of them 
exceeds 0*40/* and seven are below 0*30/1. We may therefore quite 
safely say that random samples of one hundred cysts from the same 
stool will differ from each other by more than 0*40/1 very 
rarely. If we now examine Table I we find that there are 
thirty-five different magnitudes representing the differences between 
samples of one hundred cysts from different stools of the same case 
passed on different days. Of these magnitudes sixteen exceed 0*40/1. 
Of the eight cases measured on different days, four show variations 
greater than 0*40/1. It is clear therefore that samples of one hundred 
cysts taken on different days often differ much more in average size 





6 


than samples from the same stool, and that therefore their differences 
cannot be accounted for by errors of sampling. There is thus high 
probability of accuracy for the conclusion that some infections at any 
rate are not constant, but change in average size from one day to 
another. 

It is probably worth while to add to the evidence in Case 11, the 
one showing the largest variations, by giving the results for samples 
of fifty also. The full record is then as follows : — 


Tabu: IV. 


Case 

1 

Date 1 

No. measured 

Averape diameter (/<) 

11 

17.4.1 s 

5 ° 

12-46 


3 - 5 -'* 

200 

1216 


14.5.18 

100» 

u-95 


21.5.18 

100 

1 

1214 


30*7* 18 

1 

300 

*3 33 

■ 

31.7.18 

5° 

12-92 


2.8.18 

5 ° 

13-20 


1 he full record brings out the variability in size more clearly. 

In Table I are given the dates of the periods of treatment with 
emetine. It has been suggested by Dobell and Jepps (1918) that 
emetine treatment may affect the size of the cysts in that, if the cysts 
reappear after treatment, they are liable on their reappearance to be 
larger than on normal occasions. There is only one of my results at 
all susceptible of this explanation, namely the record for Case 13 
on the first day after treatment, April 6th, 1918 (see Table I). In 
this case, however, the cysts did not disappear during treatment. It 
will be noticed, too, that other records show similar and even greater 
variations which are not susceptible of this explanation. It would 
seem therefore that there is no evidence that the variation in Case 13 
is due to the emetine treatment. 

There remains the question of the genuineness of the samples. If 
substitution of one stool for another occurred in the hospital, then the 
samples would be worthless for the purpose of this investigation. It 



7 


can only be said that the necessity of strict care in this respect was 
recognised from the outset, and that in the routine examinations in 
this laboratory extending over three years there has been almost no 
evidence that stools were sent up under the wrong name. The 
occasional possibility of this cannot be entirely excluded, but for the 
series in question at the present time the accuracy of the naming of 
• the stools was several times investigated and no reason was found to 
consider it doubtful. 

It may be remarked in passing that samples of five hundred 
cysts, such as were measured by Dobell and Jepps, ought to have an 
error of sampling considerably smaller than samples of one hundred 
cysts. Owing to the fact that the measurements are by different 
workers, my results cannot be applied in detail to their measurements. 
Since, however, my random samples of one hundred cysts from the 
same stool rarely differ by more than 0*30/1, it becomes a question 
whether 0*25/1 (the difference obtained between the first and second 
samples of Case 42 by Dobell and Jepps) is not a significant 
difference. There, is not much direct evidence as to the amount of 
variation to be expected in the average size of samples of five 
hundred cysts from the same stool. Dobell and Jepps (1918) 
measured five hundred cysts from the stool of Case E42 in saline, 
and five hundred cysts from the same stool in iodine. The average 
in saline was 12*99/1, in iodine 12*86/*, a difference of 0*13/1. Since 
iodine may be taken to have no effect on cyst size, this difference is 
probably due to sampling error. Matthews (1919) measured one 
thousand cysts of E. coli from the same stool and obtained 15*33/1 
as the average for the first five hundred and 15*27/1 for the second 
five hundred, a difference of 0*06/1. So far as they go, these numbers 
indicate that the sampling error for samples of five hundred cysts is 
about 0*10/1, and therefore that a difference of 0*25/1 is of some 
significance. 

The occurrence of change in the average size of the cysts of an 
infection obviously renders the existence of races more difficult to 
establish, but such change is in* itself quite compatible with the 
existence of races, and in fact is to be expected on that theory. 
Dobell and Jepps (1918) have themselves pointed out that, in 
mixtures of the ‘ small * with the ‘ ordinary ’ race, the cysts of the 
two component races appear in varying proportions on different 



8 


days, and thus differences in the average size of the cysts of the 
sample must necessarily be produced. In the case instanced by 
them the two component races were so far apart in size, 8 fi to 9/* 
and 1 2 /i to 14 n respectively, that the two races could easily be 
distinguished. But if races of 1 i ' 6 p and 13 3 fi exist, then no doubt 
mixtures of these will exist also. In such a mixture the two races 
might appear in varying proportions on different days, producing 
differences in the average size of the whole infection without it being 
possible to disentangle the two components. The curve of such 
mixtures would usually be unimodal and the proof of the existence 
of the two races in the mixture would be difficult, if not impossible. 

There remain, however, further tests of the validity of the theory 
of the existence of such closely neighbouring races. If the average 
sizes of all the infections measured are placed in order of magnitude, 
then if pure races are common, the averages will tend to group 
themselves around two or three nodal points corresponding to the 
two or three pure races supposed to exist. If mixtures of races are 
frequent this might not be the case, but it might* even in this event, 
be possible by constructing curves of the infections to find out 
whether the races were pure or mixed and to fix the pure races, 
and from these to disentangle the component parts of the mixtures. 
Table V shows how far this has been possible in my material. 


Table V. 


Case 

1 

No. of cysts measured 

Average size ( ji ) 

Modes of the curve 

1 

100 

11 -6 

i *'9 

2 

100 

11*6 

11*0 

3 

100 

M.7 

1 i-o and 127 

4 

100 

12-2 

* 1*9 

5 

100 

I 2’3 

n *9 

6 

100 

124 

iro and 127 

7 

100 

124 

127 

8 

100 

I2-S 

127 

9 

500 

12-6 

u*9 and 127 

10 

400 

127 

127 

11 

850 

127 

n*9 (and 127) 

12 

*75 

12-9 

127 and 13 6 

13 

200 

13-0 

127 

H 

100 

131 

136 

! 5 

178 

* 3 ** 

127 and 13-6 

16 

200 

* 3 ’* 

127 

17 

300 

13-2 

, > 3*6 

18 

200 

i 3‘5 

127 and 14*5 

19 

100 

137 

136 

20 

140 

13*8 

* 3*6 



9 


In the first place it is noticed that the average sizes of 

the twenty infections increase from i i ' 6 /i to 13*8 fi by very 

gradual steps. There are no marked gaps in the series. The 

largest gap is between Infection 3 and Infection 4, and this may 

indicate the separation out of a smaller race averaging ir6/1. 
Consideration of those infections in which fifty cysts have been 
measured goes, however, against this view. Of the thirteen 
infections in which samples of fifty to one hundred cysts have been 
measured, no less than seven show averages between 117/* and 
12*2/1, as is seen in Table VI. 

Tabit VI. 


Case 

No. of cysts measured 

Average diameter (//) 

2* 

5° 

ll *7 

22 

5° 

i*7 

23 

5° 

11-8 

24 

5° 

119 

23 

5° 

119 

26 

5° 

11*9 

27 

5° 

120 

28 

87 

12*1 

29 

5* 

12! 

30 

87 

12*2 

31 

92 

124 

| 

32 ; 

66 

127 

33 

5° 

12-8 


Though these figures are not so reliable as those from larger 
samples, yet it seems highly improbable that the measurement of 
larger samples would have removed all these infections into the 
lower or the higher of the two groups indicated in Table V. It 
seems likely, therefore, that the gap in the gradual series of averages 
in Table V is not significant. There is thus no obvious indication 
that the averages group themselves round nodal points. We must, 



therefore, pass on to consider the curves of the individual infections. 
It will be clear from what has been said that, on the hypothesis of the 
existence of further strains within the ‘ordinary* strain, (^i) those 
infections which show considerable variation in size from one day 
to another should be mixtures of races, and (2) those infections whose 
average size when well-established lies intermediate between the 
sizes of supposed pure races should be mixtures. Let us test 
these hypotheses by reference to the curves of individual infections. 
Firstly with regard to variable infections. Reference to Table I 
will show that Cases 10 and 11 had infections showing considerable 
variability. In these sufficient cysts were measured to make the 
averages and the curves very reliable. The curve for Case 10 is 
given in fig. 1, for Case 11 in fig. 2. There is in neither of them 
any clear evidence that the infection is a mixture. The curve in 
fig. 1, on the contrary, is unimodal and one of the most nearly 
symmetrical of all, showing every sign of the curve of a pure 
race. Cases 9 and 16 show very small variations such as might 
be due to errors of sampling. Their curves (Case 9 in fig. 3, 
Case 16 in fig. 4) are very similar to those in figs. 1 and 2, 
showing if anything rather less sign of purity of race than the latter. 
It may be concluded that there is no proof of the variable infections 
being mixtures. We will next consider whether those infections 
intermediate in size between supposed pure races are mixtures, and 
will examine first whether the results themselves as seen in Table V 
suggest any particular sizes as belonging to pure races. In Table V, 
column 4, are given the modes of the curves for each infection. In 
some cases the curves are bimodal, in some unimodal. The bimodal 
curves are, however, scattered quite irregularly through the series. 
The bimodal curves are more likely to represent a mixture of races 
than the unimodal, but on account of the irregularity of distribution 
of both, the table gives no evidence of the position of the 
hypothetical pure races. It remains to be seen how far the races 
suggested by Dobell and Jepps can be fitted into the facts of 
Table V. These authors have given 1 \ '6fi and 13*3/1 as the average 
diameter of two races within what has been called here the 
‘ordinary’ strain. The existence of the 1 16fi strain is not 
incompatible with the figures of Table V. Also there are sufficient 
infections with an average diameter about 13*1/4 to form some 



Number of Cysts Number of Cysts Number of C jsts Number of Cysts 


II 



10 11 12 13 14 15 16 17 


Diameter of cysts in fi . 



10 11 12 18 14 15 16 17 


Diameter of cysts in p . 

Fig. 3. Case 9. 500 cysts. Average diam. i 2 '(>fi. 







12 


confirmation of a race of about this cyst diameter. My material, 
however, provided many cases with average diameter about 12*5/1, 
i.e. about midway between n*6/* and 13*3/*, and in three of these 
cases, viz., Cases 9, 10 and 11, I measured large numbers of cysts in 
order to get a reliable curve for each infection so that it might be 
ascertained whether these infections showed any signs of being 
mixtures of the hypothetical 11*6/1 and 13*3/* races of Dobell and 
Jepps. As the curves of figs. 1, 2 and 3 show, there is no definite 
evidence in either direction. Case 10, fig. 1 (average 12*7/1), has 
a very symmetrical curve just such as would be expected from a 
pure race. Case 9, fig. 3, shows a rather less regular curve with a 
flat top, and Case 11, fig. 2, is very similar. One would *suppose, 
since cases of this average diameter occur so commonly and since 
they form fairly symmetrical curves, that a race of about 12*5/1 
existed. There is, however, similar evidence for the existence of 
races with average diameters of, say, 12*2/1 and 12*9/1, and t° 
hypothecate the existence of so many races so close to each other in 
average diameter is to render the whole theory incapable of proof.* 
Matthews (1919) has shown that in Entamoeba coli there is 
evidence of a pure race of average diameter 15 3/1 and another of 
average diameter 18*5/1, while the infections (which are common), 
having an intermediate diameter, show signs of being mixtures and 
give in almost every case a bimodal curve. The curve for the whole 
species when equal samples are taken from each infection is itself 
bimodal. The case is very different for E. histolytica . Fig. 5 
shows the curve for the ‘ordinary* strain of E. histolytica . It is a 
curve of the measurements of one thousand nine hundred cysts in 
which each of nineteen cases makes an equal contribution of one 
hundred cysts. It is unimodal and fairly symmetrical, and gives no 
evidence of any sub-division of this strain into two or more 
components. It resembles very much the curve given in my previous 
paper for the same strain, a curve which was composed of a larger 
number of smaller samples taken at random in the course of the 
routine work in this laboratory. 

• Cf. on this point Dobell and Jepps (1918), p. 340. ‘ It must be expressly noted that 

the strains of E . histolytica just described constitute merely a selection from those which we 
have studied. We have no doubt that, had it been possible for us to make an equally detailed 
study of all the cases in our series, we should have found other races possessing cysts with 
other mean diameters.’ 



13 


It seems, therefore, that by all the tests so far applied the 
existence of further strains beyond the ‘ small ’ and the ‘ ordinary * is 
not confirmed. In fact, the interesting curve for Case io shown 
in fig. i is rather strong evidence to the contrary. Here is an 
infection which shows considerable variability from one day to 
another, and moreover lies intermediately between the 11 ‘6 ft and 
*3*3/* races of Dobell and Jepps. On both counts it should be a 
mixture of races, yet its curve is unimodal and almost exactly 
symmetrical, a curve such as would be given by a pure race. The 


Fig. 5. 1900 cy6ts. Average diam. 12 ' 6 /a. 



strongest argument for the existence of further races is the 
undoubted fact that two races, the ‘ small 9 and the 4 ordinary *, do 
exist. Their existence is susceptible of proof, because their mean 
sizes are so far apart that a curve representing a mixed infection 
is certain to be bimodal. Any daily variations in size so far 
measured or any possible errors due to sampling are small as 
compared with the difference between the two races. The existence 
of the finer races is not suceptible of proof either because they do 
not exist—in which case variations between individual infections 
must receive some other explanation—or because they are so 



! 4 

numerous and so close together in average size that curves cannot 
show which infections are mixtures and which pure races. Variation 
from day to day and errors of sampling are large compared with the 
differences between the average size of these finer races.* 

Finally in Table VII and fig. 6 are given some results as to the 
prevailing size of the cysts of infections of E . histolytica among 
people from various localities. It will be seen that there is a good 
deal of evidence that in convalescent dysenteries returned to this 
country from various theatres of war, about one-third of all the 
infections of E . histolytica are those with the ‘small* race. The 
figures from Liverpool and those given by Dobell and Jepps (1917) 
agree in this, as do figures kindly supplied to me by Dr. D. L. 

Table VII. 

Size of Infections. 


ABROAD (Dysenteries) 


Cases 

Infections 

* Ordinary * 

% ‘ Small ’ 

i 

0 / 1 

0 

Authority 

i 

56 

59 

38 

6+ 

21 

* 

1 

j Mackinnon, ‘ Chronic * 
Dysenteries 

209 

225 

106 

47 

1 *9 

53 

Mackinnon (1918) 

200 

215 

140 

65 i 

75 

35 

Dobell and Jepps (1917) 

306 

325 

215 

66 

' 

1 

1 

I to J 

1 

34 

Matthews and Smith, 
convalescent dysen¬ 
teries examined at 

1 Liverpool in 1917 and 
1918 




Majority 

Minority Wenyon and O’Connor 
(' 9 > 7 ) 


NEVER ABROAD (Healthy Carriers) 


Cases 

Infections 

‘ Ordinary ’ 

0/ 

,0 

‘ Small ’ j 

0 / 

/o 

Authority 

98 

99 

8+ 

85 

15 

! 

*5 

Matthews and Smith, 
various samples of the 
home population ex¬ 
amined in 1917 and 
1918 

• 




* My measurements have not revealed a single example of the 15/1 strain of Dobell and 
Jepps, of which they appear to have had one case, and of which Wenyon and O’Connor 
record one case. It is evidently very rare. I cannot, therefore, discuss the evidence for the 
existence of this strain. This fact, however, does not appreciably affect my main conclusions. 




i5 


Mackinnon as to the incidence of such ‘ small * infections among her 
‘chronic* dysenteries.* It will be seen that, compared with this, the 
‘small* race is comparatively rare in those who have never left 
England, being present in only 15 per cent, of the ninety-nine 
E. histolytica infections found in this group. It will be noticed on 
the other hand that the ‘ small * race is remarkably prevalent among 
the two hundred and nine cases discussed by Mackinnon (.1918). 
The significance of these facts is not at all clear. A further 
characteristic of the size of infections of the ‘ ordinary * strain, in 
people never out of England is illustrated in fig. 6, where it is shown 
that the average size of one thousand one hundred and fifty-three 


Fig 6 11 53 cysts. Average diam. 12*15/4. 

°* * 16 . 1510 cysts. Average diam. 12*76/4. 



cysts from fifteent such infections in people who have never been 
abroad is 1215/4, that of one thousand five hundred and ten cysts 
from eighteent infections of the ‘ordinary* strain from those who 
have been abroad is 1276/4. The difference in the average of such 
large numbers seems significant, and thus the infections of those who 
have never been out of this country appear to be characterised not 
only by a very small proportion of the ‘ small * race, but also by a 
reduced proportion of the larger sizes of the ‘ ordinary * race. It 
may be that further measurements would tend to abolish these 

• ‘ Chronic * dysenteries were arbitrarily defined as cases that had suffered from 
dysentery, with little or no real intermission, for at least a year previous to the examination 
mad£ by Dr. Mackinnon. 

t All infections in which 50 or more cysts were measured have been included, but no case 
contributes more than 100 cysts to the curves. 




i6 

characteristics, though both the number of infections and the number 
of cysts in both categories is large. So far as the present evidence 
goes these characteristics exist, and they tend in the direction of the 
supposition that size of cyst depends to some undefinable extent 
upon locality. 

It will be seen, too, that those who have never been out of this 
country are healthy carriers, as opposed to the convalescent 
dysenteries from abroad. Thus the occurrence of a reduced 
proportion of the very small and of the very large cysts is 
characteristic of these healthy carriers. Whether there is any 
significance in this fact it is impossible on the present evidence 
to say. 


SUMMARY 

1. Evidence is given that not all infections of Entamoeba 
histolytica remain constant from one day to another in the average 
size of their cysts. 

2. While the species E. histolytica can undoubtedly be divided 
into two races characterised by smaller and larger cysts respectively, 
the existence of further races is not confirmed. 

3 Infections with E. histolytica in healthy carriers who have 
not been out of this country are characterised by a smaller proportion 
of the ‘small* race, and also by a reduced proportion of the larger 
cysts of the ‘ ordinary * race, as compared with infections from 
convalescent dysenteries from abroad. 


REFERENCES 

Dobell, C. and Jepps, M. W. (1917)- On the three common intestinal cntamocbac of man, 
and their differential diagnosis. Brit. Med. Journ. Vol. II, p. 607. 

- (1918)- A study of the diverse races of Entamoeba histolytica distinguishable from 

one another by the dimensions of their cysts. Parasitology. Vol. X, p. 320. 
Mackinnon, D. L. (1918). Notes on the intestinal protozoal infections of 1,680 men examined 
at the University War Hospital, Southampton. Lancet , Vol. CXCV, p. 386. 

Matthews, J. R. (1919). A mensurative study of the cysts of Entamoeba coli. Ann. Trop. 
Med. & Parasitol . Vol. XII, p. 259. 

Smith, A. Malins (1918). Measurements of and observations upon the cysts of Entamoeba 
histolytica and of Entamoeba coli . Ann. Trop. Med. & Parasitol. Vol. XII, p. 27. 

Wenyon, C. M. and O’Connor, F. W. (1917). Human intestinal protozoa in the Near East. 
Wellcome Bureau of Scientific Research , also Journ. R.A.M.C. Vol. XXVIII., p. 151. 



'7 


THE COURSE AND DURATION OF AN 
INFECTION WITH ENTAMOEBA COLI 

BY 

J. R. MATTHEWS, M.A. 

From the Liverpool School of Tropical Medicine 
(Received for publication 17 February , 1919) 


Little is known at present as to the length of time that infections 
with intestinal protozoa may persist, and details of the course of 
such infections have been published in only a few instances. Data 
of this kind can be obtained only by daily observation of infected 
cases over long periods of time, and opportunities for such investiga¬ 
tions seldom present themselves. Published records relate chiefly 
to infections with Entamoeba histolytica , though infections with 
E. coli and with Giardia intestinalis have also been followed for 
considerable periods. The evidence, so far as it goes, favours the 
idea that an infection, once acquired, persists for a very long time, 
if not throughout the lifetime of the host. 

Some records of examinations conducted in this laboratory show 
that several cases, infected with E. histolytica and intractable to 
treatment with emetine, have remained infected for seven to ten 
months—the time during which they were under our observation. 
It is pretty certain that these cases were infected for a considerable 
time before they reached us, and we know that some of them 
remained infected for some time after their departure, until we 
ultimately lost trace of the cases. But infections of one or two years* 
duration are short compared with those given for three cases 
described by Dobell and Stevenson (1918). These authors, from the 
evidence at their disposal, conclude ‘ that an infection with 
E. histolytica may persist certainly for sixteen, and probably for as 
long as thirty-four years.* 

It is of interest, therefore, to enquire if the harmless intestinal 
protozoa behave similarly in this respect. It has been shown in a 



i8 


recent paper (Matthews and Smith (1919)) that infections with 
Giardia intestinalis and with E. coli are more prevalent among 
children than among adults.* The authors suggested an explana¬ 
tion of these observations on the supposition that infections had been 
lost among the adults. The evidence, however, was indirect, and 
the purpose of the present note is to record certain observations which 
seem to furnish fairly satisfactory reasons for believing that an 
infection with E. coli , at all events, may disappear from the intestine 
in the course of.time. 

The individual whose infection has been followed was first 
examined for intestinal protozoa at approximately fortnightly 
intervals during the first three months of 1916. These examinations 
were negative. In the first week of August, 1916, further 
observations were made, when cysts of E. coli were detected in the 
stools. It cannot be definitely concluded that the infection was 
acquired just previous to that date, but since the early examinations 
were negative and since the person has from the beginning of 1916 
been engaged on the protozoological examination of stools, it seems 
not unlikely that infection occurred in the laboratory between 
March and August, 1916. In November, 1916, a series of daily 
examinations was commenced, and was continued for three months. 
The main object of these examinations was to obtain data which 
would give a rough idea of the course of the infection over a 
considerable period of time. The method adopted was to count the 
number of cysts contained in six cover-slip preparations made from 
different parts of the stool. The accompanying chart (fig. 1) shows 
the average number of cysts for each preparation for a period of two 
months, t The method employed cannot be expected to give more 
than a rough idea of the course of the infection. Cysts of E. coli 
are usually irregularly distributed in the stool whether they are 
numerous or not. On one occasion six preparations contained 
respectively 76, 43, 39, 197, 143 and 79 cysts. Again, on a day when 
the cysts were much less numerous, six preparations contained 
respectively the following numbers of cysts—2, 7, 1,0, 1, 4. The 


• The figure* were : Among children— G. intestinalis , 14*1 per cent.; E. coli , 111 per cent. 
Among adult*—6-o per cent, and 67 per cent., respectively. 

t It should be mentioned that there wa* only one stool daily, although on two occasions, 
December 10th and January nth no stools were passed. 



19 



«j«Xj JO Jaquin^ 


20 


preparations each day were made as uniformly as possible, though 
slight variations in this respect doubtless occurred from day to day. 
It is believed, however, in spite of the defects of the method, that 
the main features of the course of the infection are brought out, and 
the rise and fall in the number of cysts produced daily is a feature 
which must be familiar to anyone who has followed infected cases 
for any length of time. It is this feature that gives in a very general 
sense an indication of periodicity in cyst production (cf. Porter’s 
(1916) results for Giardia). An examination of the chart shows 
that the maximal values occur at intervals of 8, 8, 5, 8, 8, 8 and 7 
days respectively, and the minimal values at intervals of 8, 8, 9, 4, 
9, 10 and 6 days respectively. There is thus no conclusive evidence 
of regular periodicity in cyst production. It should be remembered, 
also, that we have here the data for a single case only, and it is 
unlikely that all infections with E. coli would behave in an exactly 
similar way. 

Daily observation of the case was discontinued in February, 1917, 
only occasional examinations being made until July, 1917. Without 
exception these were positive. Thereafter, twelve months elapsed 
before the case was again examined. Several observations in July, 
1918, gave negative results, and in view of the long succession of 
positive findings in the early history of the case, it seemed of interest 
to discover how long the negative period that had been encountered 
would extend. From 18th July, 1918, to 15th February, 1919, eighty 
negative examinations have been made, or an average of three 
examinations every eight days during a period of seven months. 
There now arises the difficult question of deciding whether the* 
infection has entirely disappeared or whether a very long negative 
period has set in. Many cases might be cited from records obtained 
in this laboratory showing long intervals during which no cysts were 
found in the stools, although the cases were known to be infected. 
All these cases, however, are complicated by the fact that they were 
also infected with E. histolytica and had received treatment with 
emetine. It is not certain how far this treatment had an inhibitory 
effect on the concurrent infection with E. coli. Again, in instances 
where E. coli is recorded at a late examination (e.g. in one case at 
the fifty-fifth examination after three months* observation) there is 
no certain proof that the man had not recently become infected, 



21 


i.c. not long before the positive finding was recorded. Finally, there 
are cases where E. coli has been found only once in a long series of 
examinations (e.g. one positive record out of sixty-five examinations 
extending over a period of five months). In such cases, there is no 
absolute certainty that the sample submitted, on the occasion when 
E. coli was found, was genuine. These criticisms, however, do not 
apply to the case which forms the subject of this note. Emetine 
was not administered in any form, and there was no doubt about the 
genuineness of the samples of stools examined. That the case has 
remained negative for seven months seems to indicate without much 
doubt, although not with absolute certainty, that the infection has 
disappeared entirely. It may be affirmed, anyhow, that should 
E. colt again be found in the same case, there will be no means of 
proving that re-infection has not occurred. 

The facts of the present case may thus be summarised. Infection 
was known to exist for a year—from August, 1916, to July, 1917. 
A year elapsed without examination. Since July, 1918, frequent 
negative examinations have been recorded, and it is believed that 
the infection has been lost. 

We have no means of deciding whether the course and the 
duration of the E . coli infection here described are normal (see 
Wenyon and O’Connor, 1917, p. 71). It is doubtless true that many 
cases remain infected for much longer than a year. James (1914) 
has mentioned the case of a negro known to have had an infection 
with E. coli lasting at least six years. We do not know, however, 
how often infections of short duration occur. If they are at all 
common, they may give rise to a factor which should be borne in 
mind when dealing with the comparative incidence of infection 
among different sections of any population examined for intestinal 
protozoa. As has already been mentioned, we have recorded 
significant differences in the incidence of infection with Giardia 
intestinalis and Entamoeba coli among Liverpool children and 
adults. 

There does not appear to be any reason why E. coli should not 
disappear spontaneously. All the amoebae in the intestine may 
perish in some way, or they may all encyst simultaneously and all be 
evacuated in the encysted form. In either event the infection would 
be lost. 



22 


REFERENCES 


Dobell and Stevenson (19x8). A note on the Duration of Infections with Entamoeba histolytica. 
Tram. Trop. Med. & Hyg. Vol. XI, pp. 168-175. 

James (1914). A Study of the Entamoebae of Man in the Panama Canal Zone. Atm. Trop. 
Med. Parasitol . Vol. VIII, pp. 133-320. 

Matthews and Smith (1919). The Spread and Incidence of Intestinal Protozoal Infections 
in the Population of Great Britain. III. Children. Ann. Trop. Med. & Parasitol. 
Vol. XII, pp. 363-371. 

Porter (1916). An enumerative study of the cysts of Ciardia (Lam Mi a ) intestinalis in human 
dysenteric faeces. Lancet, 10th June, pp. 1166-1169. 

Wenyon and O’Connor (1917). Human Intestinal Protozoa in the Near East. London. Wellcome 
Bureau of Scientific Research. 



*3 


TWO PARASITES OF NAJA 
NIGRIC OLLIS 

BY 

J. W. S. MACFIE 
(Received for publication 16 October , 1918) 

(Plate I) 

I 

TRYPANOSOMA VOLTARIAE, ap. n. 

(Plate I, figs. 1 to 4) 

This trypanosome was found in the blood of a snake in the 
Gold Coast Colony, West Africa. The host was identified as 
Naja nigricollis. 

The infection was a very scanty one, only about half a dozen 
trypanosomes being found in each of the blood-films obtained. The 
parasites were not seen alive. The description is therefore based 
entirely on the appearance of the parasite in blood-films stained with 
Leishman's stain. The trypanosome was monomorphic; all the 
specimens examined having a similar shape and being of about the 
same size. 

All the trypanosomes found in the films were looped, and had 
the posterior extremity bent at a right angle to the rest of the body 
(figs. 1 to 4). The posterior end of the body extended for a 
considerable distance beyond the blepharoplast as a snout, which was 
always more or less folded. The body, which stained a blue colour, 
was compact posteriorly, but anteriorly frayed out into a number 
of striae which blended with the undulating membrane. The 
blepharoplast was small, rounded or oval, and compact, and was 
situated at some distance posterior to the nucleus. The nucleus was 
oval, measured about 3 p long and 2 p broad, and lay on the same 
side of the body as the undulating membrane at a point slightly 
posterior to the middle. The undulating membrane was well 
developed and arranged in ample folds along the external border 



H 


of the loop formed by the body of the trypanosome. The flagellum 
terminated anteriorly in a free portion. The average measurements 
of five parasites were as follows:—Total length, 50 fi; breadth (not 
including the undulating membrane) at the level of the nucleus, 4 a* ; 
posterior extremity to blepharoplast, 17A 4 ; blepharoplast to nucleus, 
4 n; free portion of the flagellum, 8 a*. 

This trypanosome, so far as I am able to ascertain, differs from 
any of the species hitherto found in snakes. In some respects, 
however, it resembles the small forms of T. primeti , a species found 
in the blood of water snakes in Tonkin, and described by Mathis and 
Leger (1909). These forms of T. primeti appear to be somewhat 
longer (57/1) and broader (7/1) than the trypanosome of Naja 
nigricollis described above, the distance from the blepharoplast to 
the nucleus (2/1) is shorter, and the free portion of the flagellum (13/1) 
is longer. Moreover, with the small forms are associated others 
about twice their size, both being comprised in the species T . primeti 
by Mathis and Leger. In the snake examined at Accra no trypano¬ 
somes were found resembling the large forms of T. primeti. Too 
much importance must not be attached to the small differences in 
measurements, because in the case of the trypanosome of Naja 
nigricollis only a very few individuals were examined, and in the case 
of T . primeti nothing is said about the range of variation. Mathis 
and Leger do not mention any forms intermediate in size between 
/ the large and small forms of T. primeti y but they appear to have been 
satisfied that only a single species of trypanosome was represented, 
and considered that the small forms were the young forms. 

It should be mentioned that Wenyon (1908) has described a 
trypanosome, T. najae y found in Naja nigricollis in the Sudan, but 
as he saw it only in fresh blood, and as his drawings made from 
living specimens are mere sketches, it is not possible to compare it 
with the parasite found in the Gold Coast. For the latter parasite 
I propose the name Trypanosoma voltariae . 

REFERENCE 

Mathis, C., and Leger, M. (1909.) Sur un Nouveau Trypanosome des Serpents du Tonkin. 
Comptes rend us de la Soc. dc Biol ., LXVIT, pp. 572-574. 

Wenyon, C. M. (1908). Report of Travelling Pathologist and Protozoologist. Third 
Report of the IVell come Research Laboratories at the Cordon College , Khartoum , 
pp. 142-141- 



25 


II 

PLASMODIUM MESNIU, Bouet 

(Plate I, figs. 5 to 24) 

Pigmented parasites of the red blood corpusdes have been 
described by several authors as occurring in snakes, but hitherto 
only gametocyte forms have been seen in the blood, and these appear 
to have belonged to but a single species. The name Plasmodium 
mesnili , given by Bouet (1909) to the species described by him, has 
therefore been applied generally to these parasites. 

In a specimen of Naja nigricollis killed in 1916 at Accra in the 
Gold Coast, West Africa, the blood was found to be heavily infected 
with pigmented parasites, and as certain stages of the life-history 
were exemplified which have not previously been described, a brief 
account of them will be given. I have to thank Dr. A. Ingram for 
very kindly sending me the materials from this snake, namely two 
blood-films stained with Leishman's stain. 

FORMS SEEN IN THE BLOOD 

A considerable proportion of the erythrocytes of the snake were 
infected by parasites, many of them doubly, and some trebly. The 
most conspicuous forms were the gametocytes, but there were also a 
great many amoeboid forms, and a few which appeared to be under¬ 
going schizogony. A few parasites, both young trophozoites or 
merozoites and rounded-up gametocytes, were seen free, but it is 
possible that this was the result of some post-mortem change or of 
mechanical injury to the corpuscles in the act of making the films. 
The infected erythrocytes stained similarly to the uninfected, 
and showed neither polychromatophilic changes nor stipplings 
(excepting the macules referred to below). Pigmented leucocytes 
were very abundant. As the trophozoites contained very little 
pigment and the gametocytes a large quantity, it may be surmised 
that much of the pigment in the leucocytes was derived from the 
latter after their death and disintegration. 

In addition to the pigmented parasites there were a few haemo- 
gregarines similar to those figured by Wenyon (1908) as occurring in 
Naja ntgricollts . 



26 


Schizogony. The youngest forms were usually situated at the 
ends of the corpuscles, and were oval or slightly irregular bodies 
with a small mass of chromatin at one pole. Somewhat larger 
forms were frequently ring shaped with a large central vacuole. 
Older parasites were often very irregular in shape, their cytoplasm 
thrust out as delicate pseudopodia, indicating a considerable degree 
of amoeboid activity (figs. 6 to n). The trophozoites stained a 
pale blue colour. When fully grown they measured about $/* in 
length. Pigment in the form of rather fine granules of a brown 
colour was sometimes present, but was generally scanty or 
inconspicuous and often entirely absent. The trophozoites did not 
cause any marked deformity of the erythrocytes or displacement of 
the nucleus. 

Two or more trophozoites were frequently present in the same 
erythrocyte. In some such cases the cytoplasm of two parasites 
appeared to have fused, giving rise to a form suggesting schizogonic 
multiplication (figs. 9 and 10), but differing from that which I believe 
to be the true multiplication form (fig. 14). Such forms are indeed 
the very reverse of division forms, being produced by the conjunction 
of separate individuals. 

Schizogony appeared to take place by the nucleus dividing into 
two with the formation of a simple dumb-bell figure, the formation 
of a binucleate parasite, and the subsequent separation of this into 
two daughter cells of about the same size (fig. 14). No indication 
of the formation of more than two merozoites was seen. The 
number of parasites undergoing schizogony in the blood was very 
small, whereas the number of trophozoites, both young and further 
developed forms, was large. It is possible therefore that schizogony 
occurs mainly in the organs. 

GAMETOCYTES. The gametocytes were of two distinct types, the 
cytoplasm of the one staining a deep blue with Leishman’s stain, 
that of the other remaining almost uncoloured. The former are 
considered to be the females, and the latter the males. The male 
and female gametocytes were present in about equal numbers: 
105 male and 95 female forms occurring in a series of 200 consecutive 
individuals counted. Whilst still quite small, the male and female 
gametocytes were readily recognised and easily distinguished from 
one another by their staining reactions. The gametocytes when 



27 


young lay usually at the ends of the corpuscles (fig. 20), when a 
little older they lay laterally, and eventually looped round the nucleus 
of the corpuscle, sometimes surrounding it completely. Many 
corpuscles contained two gametocytes of either the same or of 
different sexes (figs. 18, 19, 23). As a rule, a single gametocyte of 
either sex, even when fully grown, produced but little distortion of 
the erythrocyte, but considerable enlargement of the cell and 
displacement of its nucleus was caused by double or multiple infec¬ 
tions, and a similar result was caused by a single gametocyte if it 
developed at the end of the corpuscle instead of growing round the 
nucleus in the usual halter-like manner. The gametocytes contained 
a large quantity of dark brown pigment in the form of coarse grains. 

Both male and -female gametocytes were occasionally seen which 
had two nuclei (fig. 22), but as they were of large size, caused 
distortion of the erythrocyte, and often showed two vacuoles, it is 
probable that they were in reality each compounded of two parasites 
lying side by side whose individual outlines were invisible. This 
interpretation is in agreement with that given by Woodcock (1911) 
of similar forms of Halteridium frangillae observed by him in the 
blood of a chaffinch. The gametocytes of Haemocystidium sitnondi 
described by Dobell (1911) as showing nuclear division are considered 
by Woodcock to be subject to the same explanation. 

The male gametocyte . The male gametocyte stained very 
faintly with Leishman’s stain. When fully developed it measured 
about IS/* by 5A*. It contained a large amount of pigment in rather 
coarse grains scattered irregularly about its substance (fig. 16), but 
sometimes collected into clumps at the extremities (fig. 19). The 
nucleus consisted usually of a group of coarse chromatin granules 
arranged in a rather compact mass near the middle of the parasite. 
In some specimens, however, the nuclear chromatin was more 
abundant, forming a band across the middle of the parasite (fig. 16) 
or a large central area lightly dotted with fine granules (fig. 17). 
The male gametocyte frequently showed a large rounded vacuole 
(fig. 15), which was, however, inconspicuous in the preparations 
studied owing to the very faint staining of the parasite with 
Leishman’s stain. 

The female gametocyte. The female gametocyte stained a deep 
blue colour with Leishman’s stain. When fully developed it 



28 


measured about 16 /i by 6 p. The female gametocyte contained a 
large quantity of pigment in coarse granules of a dark brown colour. 
The nucleus was indistinct in most specimens, but consisted of 
chromatin granules arranged so as to form an oval or more elongated 
body about the middle of the parasite. A rounded vacuole was 
present in many of the female gametocytes, and was a conspicuous 
object by contrast with the deep blue colour of the cytoplasm 
surrounding it. Sometimes two vacuoles were seen in what appeared 
to be a single gametocyte, but this was probably to be explained by 
there being two gametocytes lying side by side in the same cell. A 
few of the erythrocytes containing female gametocytes showed red 
macules (fig. 24) of a characteristic scarlet colour similar to those 
described by Wenyon (1908). 


NOMENCLATURE OF THE PARASITE 

So far as the gametocytes are concerned, the parasite which has 
just been described resembles that observed by Wenyon (1909) in 
the Sudan and originally called by him Haemocystidium najae , a 
name which he subsequently (1915) admitted to be a synonym for 
Plasmodium mesnili . These forms very closely resemble Haerno- 
proleus 9 and, in fact, Franca (1917) in his recent work ‘Sur la 
classification des hemosporidies ' has included them in this Genus 
under the name Haemoproteus najae. The occurrence in the blood 
of other stages of the parasite besides the gametocytes, however, 
renders the retention of this organism m the Genus Haemoproteus 
inadmissible. 

Wenyon (1915) has proposed to group the pigmented parasites 
of cold-blooded animals, which have been variously named 
Haemoproteus , Haemocystidium , Haemamoeba , and Plasmodium , 
into two genera; ‘firstly, the genus Plasmodium, which includes 
parasites which go through the schizogony stage in the red blood 
corpuscles and produce gametocytes in these cells too; secondly, the 
genus Haemoproteus, including parasites which do not reproduce by 
schizogony in the red blood corpuscles, which cells only contain the 
gametes in varying stages of growth.' In accordance with this 



29 


classification, and because at that time only gametocytes had been 
found in the blood, Wenyon placed the pigmented parasite of the 
erythrocytes of snakes in the Genus Haemoproteus , but the descrip¬ 
tion given above shows that it should more properly be included in 
the Genus Plasmodium. 


REFERENCES 


Bouet, G. (1909). Stir deux himocytozoaires pigmentis des reptiles. Comp. rend. Soc. Biol . 
Vol. LXVI, pp. 43-45. (Jan. 9, 1909). 

Dobell, C. (1911). Contributions to the Life-History of Haemocystidium simondi , Castellani 
and Willey. Festschrift zum Secbzigsten Geburtstag Richard Herttcig , Gustav Fischer, 
Jena, Bd. I, pp. 123-132. 

Franca, C. (1917). Sur la classification des hiraosporidies. Jornal de Sciencas Matemdticas , 
Fisicas e Naturais , 3rd series, No. 1. 

Wenyon, C. M. (1908). Report of Travelling Pathologist and Protozoologist. Third Report 
of the Wellcome Research Laboratories at the Gordon College , Khartoum , pp. 150-152. 

-(1915). The Pigmented Parasites of Cold-blooded Animals, with some Notes on a 

Plasmodium of the Trinidad Iguana. Journ. Trop. Med. & tfyg., Vol. XVIII, 
pp. 133-140. 

Woodcock, H. M. (1911). On an unusual condition observed in Halteridium. Zoologiscbe 
Anzeiger (Leipzig), Bd. XXXVIII, pp. 465-471. 



30 


EXPLANATION OF PLATE 

Two parasites of Naja nigricollis . Drawings by Miss M. Rhodes 
from specimens stained with Leishman’s stain. 

Figs. I to 4. Trypanosoma voltariae t sp. n. x 1500. 

Figs 5 to 24. Plasmodium mesnili, Bouet. x 1500. Figs. 5 

to 13, various forms of trophozoites; figs. 6 to 11, showing 
amoeboid forms of the parasite. Fig. 14, dividing form. 
Figs. 15 to 18, male gametocytes. Fig. 19, double infec¬ 
tion with a male and a female gametocyte. Figs. 20 
to 24, female gametocytes; fig. 20, two immature female 
gametocytes in a single erythrocyte; fig. 21, parasite 
encircling the nucleus of the erythrocyte; figs. 22 and 23, 
double infections; fig. 24, erythrocyte invaded by a 
female gametocyte, and showing two red macules. 



Annals Trof. Med. 6 » Parasitol,, Vol. Xlll 


PLATE I 


1 


2 


3 


4 



C. Tinlmg & Co. Ltd., Imp. 




31 


ON THE GENITAL ARMATURE OF THE 
FEMALE TSETSE-FLIES (GLOSSINA) 

BY 

ALWEN M. EVANS, M.Sc. 

(Received for publication 13 March , 1919) 

INTRODUCTION 

It is well known that before 1911 the classification of the genus 
Glossina was based on highly unsatisfactory characters. Conse¬ 
quently the determination of specimens was a matter of great 
uncertainty, and this difficulty still exists in a large measure as far 
as the females of Glossina are concerned. The present paper is a 
record of investigations, the object of which was to discover 
characters of systematic value in the armature of these latter. In 
the two more specialised groups of which G. palpalis and 
G. morsitans were taken as types, the enquiry met with slight success 
in so far as the differentiation of the individual species included in 
these groups is concerned, but in the Fusca group the armature of 
each species was found to exhibit at least one distinctive character, 
and it has been discovered that six of the members of this latter 
group possess internal chitinous structures, which are specifically 
highly characteristic. 

A fact of systematic interest which has come to light is that the 
armature of the females falls into three clearly defined structural 
types which correspond to the three groups into which Glossina is 
divided on the basis of the male armature by Newstead (1911). 

I wish to thank Professor Newstead, F.R.S., to whom I am 
• indebted for the indispensable assistance which he has most kindly 
rendered to the furtherance of this investigation. 

GENERAL ACCOUNT OF THE MORPHOLOGY OF THE 
FEMALE ARMATURE 

The external armature of the female Glossina brevipalpis } 
Newst., is briefly described by Stuhlmann under the name 
of G. fusca , Walker (1907, pp. 57, 58). He refers to a ‘hufeisen- 
formiger Chitinkorper * which, he suggests, may represent the 



32 


IX° segment, and he states that within it lies the genital opening, 
below which is the anus, separated from the former by a very 
small plate Were this the case, then the relative position of 
the genital and anal apertures in the females of Glossina would 
be the reverse of that which holds in other members of the Diptera. 
It is, however, a simple matter to demonstrate by dissection (fig. i, A) 



Fig. i. A. Glossina fusca : 9- Diagrammatic representation of external armature 

together with extended uterus after maceration in KOH. Ventral aspect. a,p . 9 anal plate ; 
c.p process of signum ; cb chorion ; d.p. y dorsal plate ; g/., genital fossae ; o.d oviduct; 
p. y petaloid marking on signum ; s. y signum; sp. y spermatheca; st. vii, seventh stemite \ 
»/., uterus j v., vulva j y and z. } bristles inserted into vulva and anus respectively. 

B. Glossina fuscipUuris : lateral aspect. 








33 


that the opening between the arms of the ‘ horseshoe ’ (Chitin- 
hufeisens) leads directly into the rectum, and that the vulva (v.) is 
a broad slit-like opening between the hinder border of the VII 0 
stemite and the base of the small piece of chitin (st.p.) which 
separates the apertures. Thus the genital orifice, as in other 
Diptera, lies ventral to, not dorsal to the anus. We may now 
proceed to a comparison between the external armature of G. fusca 
and that of the blow fly, CaUiphora erythrocepJiala. In the latter 
case the ovipositor bears, terminally, four projecting plates, which 
surround the anal orifice, and are respectively referred to by Lowne 
(1893-95) as the ‘sternal plate,’ median and ventral; the ‘dorsal 
scale,’ median and dorsal; and a pair of ‘lateral scales.’ The 
vulva ( v .) is a horizontal slit between the base of the sternal plate 
and the border of the VII° stemite. We may therefore conclude 
that the small triangular plate ( st. p.) in G. fusca is the homologue 
of the sternal plate of C. erythrocepkala; this name will be used in 
describing it. The anal scales of the blow-fly which lie lateral to 
the anus are undoubtedly represented in G. fusca by the ‘Chitin- 
hufeisens,’ which, when examined microscopically, is seen to consist 
of a pair of projecting, slightly chitinised, scale-like plates bearing 
flexible spines, and here referred to as the ‘ anal plates ’ (a. p.). In 
G. fusca the median dorsal scale of the blow-fly is apparently 
unrepresented, but dorsal and external to the anal plates are paired, 
elongate, chitinisations of the integument, the dorsal plates id. p.), 
which, as Stuhlmann (l. c.) suggests in the case of G. brevipalpis, are 
probably the tergites of the VI 11 ° segment. It is the form of these 
plates which is the most variable feature of the external armature of 
the Fusca group of Glossina. 

The position occupied by the several parts of the armature in 
relation to the rest of the abdomen (in G. palpalis ) is indicated in 
fig. 2. The plates are borne by the terminal wall of the abdomen, 
a vertical membrane stretched between the arched posterior edge of 
the VIII° tergite and the posterior margin of the VII 0 stemite, from 
which latter it is separated by the slit-like opening, the vulva. 

The internal armature. In fig. 1, A, a bristle (y) is inserted 
through the vulva into the uterus, which in G. fusca and 
other members of this group bears at the anterior extremity 
of its dorsal wall the structure g. f. This is a thick semi- 



34 


transparent mass of tissue which persists after fifteen minutes* 
maceration in boiling KOH. As no reference could be found to 
such a structure in the literature on Glosstna , the vagina of Calliphora 
erythrocephala was examined, and it was found that the organ 
situated in the dorsal wall, and described by Lowne (1893-95) as 
the ‘genital fossae/ is of similar character to that here referred to 
as g. /. in G. fusca. Further, there occur on the internal surface of 
the genital fossae in C. erythrocephala two paired curved plates of 



Fig. 2. Glosstna palpalis : J. Terminal segments of abdomen, ventro-lateral aspect. 
s. vi, spiracle of sixth segment; ster. vii, seventh stemite. 

yellow chitin, which, though figured (Plate XLVII), are not named 
by Lowne (1893-95). They are represented in typical G. fusca 
(fig. 4) by a peculiar symmetrical structure composed chiefly of a pair 
of darkly chitinous, hollow, sub-conical elements (fig. 4, e), which 
will be referred to provisionally as the signum , a term applied by 
Pierce (1914) to ‘an internal armature of the bursa* in certain 
Lepidoptera. Although, as Lowne (1893-95, p. 577) states that the 
‘ young uterus * is termed ‘ the bursa,* there seem to be some grounds 
for regarding these structures as homologous, I do not commit myself 
to this view since I have not had the opportunity of examining fresh 
material of female Glosstna , but merely use Pierce’s term in the sense 
that it refers to internal armature. 








35 


TECHNIQUE 

The method employed is as follows: The terminal portion of 
the abdomen is cut off and placed in a test-tube containing io per 
cent. KOH. The latter is allowed to boil in a water bath for fifteen 
minutes, after which the specimen is well washed in water and the 
dissolved tissues expressed. The best result will be obtained if the 
abdominal wall is slit laterally, almost to the extremity, so that, in 
mounting, the specimen can be spread out with the plates of the 
armature lying neither above nor below any other portion of the 
integument. Unless a permanent preparation is required, the 
specimen may be mounted in glycerine for examination. In the 
former case it is dehydrated, cleared in oil of cloves, and mounted 
in Canada balsam. 

To dissect out the signum, in the Fuse a group remove the last 
four segments, and, if the presence of a larva is indicated, the whole 
abdomen. After the specimen has been macerated for fifteen 
minutes in KOH, and washed in water, the abdominal walls are slit 
and the contents teased out with a pair of needles. With the aid 
of a lens a pair of sub-spherical yellow bodies will be detected; these 
are the spermathecae, they lie in close proximity to the distal end 
of the uterus, to which they are connected by their much convoluted 
ducts. If fossae are present they will appear as a gelatinous mass 
occupying the extremity of the uterus, and on the surface will be 
seen a dark red-brown or ochraceous, symmetrical structure—the 
signum. When found, the genital fossae with the signum should 
be isolated and mounted in glycerine, or after dehydration in 
Canada balsam. 


Table for the Determination of Groups 

i. Dorsal Plates absent . Morsttans Group (III) 

Dorsal Plates present .2 

Medio-dorsal Plate absent... 

Medio-dorsal Plate present 


2. 


... Fusca Group (I) 

... Palpalis Group (II) 



36 


Table for the Determination of Species of Fusca Group 


1. Signum of uterus absent. 

Signum of uterus present. 

2. Hamate sclerites at base of sternal plate 
No hamate sclerites at base of sternal plate 

3. Dorsal plates narrowly crescentic . 

Dorsal plates not narrowly crescentic . 

4. Signum markedly elongate and strongly flexed ... 
Signum not markedly elongate or strongly flexed 

5. Signum consisting of two narrow, pale, parallel 

strips. 

Signum not consisting of two narrow, pale, parallel 
strips. 

6. Signum chiefly composed of a pair of conical or 

hemispherical lobes. 

Signum composed of two long, convergent bands 
separated by a deep median cleft . 

7. Dorsal plates of great extent; signum consisting 

of two entirely separate, paired sclerites ... 

8. Dorsal plates sub-triangular and not very large ; 

signum consisting of a single plate. 


2 

3 

G. brevipalpis 
G. longipennts 

4 
7 

G. fuscipleuris 

5 

G. nigrofusca 

6 

G. fusca 
G. medicorum 
G. sever ini 
G. tabaniformis 


GROUP I. Fusca Group (figs. 3-14). External armature con¬ 
sisting of five plates—one pair dorsal^ one pair lateral, and 
a single median sternal one; medio-dorsal plate absent . 
Signum generally well developed . 

External armature . Much greater in extent than in either of the 
other two groups, the dimensions varying between about n x 
o*8 mm. and o*8 x 07 mm. The dorsal plates ( d . p.) are usually 
surrounded by a considerable area of unchitinised, membraneous, 
integument which separates them from the anal plates ( a . p.'). 
The outline of the dorsal plates is generally elongated; their long 
axes lying obliquely and approaching one another dorsally. They 
always bear stout, black, spines. The anal plates (a. p.) usually 
project almost vertically from the posterior surface of the abdomen, 
to which they are only attached basally. They can be readily 
turned outwards with dissecting needles. 

Internal armature . In the species G . fusca , G. tabaniformis, 




37 


G. fuscipleuris, G. medicorum, G. nigrofusca and G. severini, there 
occurs associated with the internal genitalia the peculiar chitinous 
structure here referred to as the signum. This is always of 
symmetrical form, and affords, apart from the external armature, 
important morphological characters for the distinction of species. 

Glossina fusca, Walker 

External armature of the female (fig. 3). The dorsal plates 
{d. p.) are sub-crescenfic, the ends being more or less narrowly 
rounded or truncated. The greatest width is generally not more 
than one-fourth of the total length, and stiff spines are borne on 



Fxo. 3. Glossina jusca : 9 * External armature, x c. 80. Lettering at in Fig. i. 


the entire surface. The anal plates (<z. p.) project almost vertically 
one at each side of the sternal plate, and, when mounted in dorso- 
ventral aspect (fig. 3), each presents a narrow crescentic outline. 
When viewed in a horizontal position they are seen to have a 
somewhat rectangular form (fig. 1, a. p.) They bear setae which are 
much smaller, finer, and more flexible than those borne on the dorsal 
plate. 






3 » 


Signum of (he uterus (fig. 4). The genital fossae are oblong-ovate 
in outline, the long axis being transverse. The signum of a typical 
G. fuse a is composed mainly of a pair oi sub-conical, hollow and 
highly chitinous lobes (*.), the colour varying from dark mahogany- 
brown to black. The inner surface of each lobe on either side of 
the median line, in front usually bears more or less strongly defined 
ridges. In dorso-ventral view a crescentic sclerite ( cr .), which may 



Fig. 4. Glossina jusca : Signum, x r. 1 55. cr ., crescentic sclerite; e., main elements; 

p., posterior plate ; other lettering as in Fig. 1. 


appear densely black, bounds or overlies the posterior or postero¬ 
lateral border of each lobe. In lateral view this is seen to be in 
reality a curved plate of chitin projecting from the base of the lobe. 
At its upper extremity it is sharply bent, so that the posterior 
portion appears in surface view as a pale ochraceous plate (fig. 4, p.) 
completing the signum behind. 

Type of female armature taken at Tainsu, Wenchi, N. Ashanti, 
24.4.10 (Dr. A. Kinghorn). Five other typical examples were from 
the following localities:—Sunyani, Ashanti (2), 18.10.13 (Dr. F. H. 




39 


Storey); Volta River, Kpong, Gold Coast,. 9.1918 (Dr. P. D. 
Oakley); and Tekiman Territory, Ashanti, 4.1912 (Dr. T. E. Fell). 

The nine examples of G. fusca from the Katanga district of the 
Congo Free State taken by Dr. J. Schwetz were found to exhibit a 
marked deviation from the form of signum described above. The 
main elements or lobes (fig. 1, A, g.) are sub-hemispherical, semi¬ 
transparent, and of ochraceous colour; they bear in front a median 
collar-like projection (a. p.), and in addition a pair of distinct dorsal 
thickenings gives rise to the conspicuous, petaloid marking (J >.). 
In two examples from the Belgian Congo (ex. coll. Mus6e Royal 
d’Histoire Naturelle de Belgique), the internal wall was partially 
unchitinised and the condition was somewhat intermediate between 
the typical form and that of the Katanga specimens, though it more 
closely resembled the former. The signum of specimens from 
Buamba Forest, Semliki Valley, 2,300-2,805 feet, Uganda Pro¬ 
tectorate, 3-7 November, 1911, was largely unchitinised posteriorly, 
and anteriorly presented features in common with the specimens 
from Katanga; the median anterior projection and petaloid markings 
were conspicuously present. 


Glossina nigrofusca, Newstead 



External armature of the female. Resembles that of G. fusca. 
Signum of the uterus (fig. 5). This is slighlty developed, 
consisting solely of a pair of small elongate strips of pale yellow 



4 <> 


chitin. The surrounding membrane is thrown into innumerable 
folds of intricate character, a pair of which traverses the bars of the 
signum, giving rise to a fold (fig. 5, /.) near the middle distance. 

Type of female armature taken at Sunyani, Ashanti, 9.6.13 
(Dr. F. H. Storey), other examples at Fiapri, Ashanti, Gold Coast 
(Dr. F. H. Storey) received 17.6.13; and Boonso, Birrim District, 
Gold Coast, 9.1912 (Dr. David Duff). The form of signum here 
described* was met with also in two specimens from Tain River, 
Nsoko, W. Ashanti (Dr. A. Kinghom), one taken 7.5.10, the other 
21.5. io. The antennae were, however, not specifically identical 
with those of G . nigrofusca , but resembled those of G. fusca, , while 
the thoracic markings were paler than in typical forms of either of 
these species. A parallel case has been found in the males; a 
specimen from Ashanti (Dr. A. Kinghom) having internal armature 
typical of G. nigrofusca , antennae resembling those of G . fusca , and 
pale thoracic markings. 

Glossina fuscipleuris , Austen 

External armature of the female. Differs in no essential respects 
from that of G . fusca. 

Signum of the uterus (fig. 6). A narrowly elongate structure 
the length about four times the greatest width. It is strongly 
flexed in the middle of its length (fig. 6, c.), the two limbs 
being almost at right angles. Fig 6 , A, shews the signum in dorso- 
ventral aspect, the internal surface uppermost, having been 
straightened by pressure. In this position it appears as a ligulate, 
strongly ochraceous, lamina (/.) of somewhat irregular outline with 
a more or less deep bifurcation distally. Proximally a stem-like 
portion projects backwards carrying a pair of small lateral 
expansions (t. e.). This projecting process consists of a pair of very 
thick strongly chitinous ridges (/. p .), which form a prominent keel 
beneath the posterior surface of the lamina (fig. 6 , C.). In the middle 
length of the latter arise at the borders a pair of band-like anterior 
thickenings (/. a .), which run forward supporting the lamina and 
converging distally. Medianly the chitin is thin, and may give 
place to one, two or more small apertures. When mounted without 
manipulation the signum tends to lie in a lateral position as shown 



4 * 


in fig. 6, C. Viewed thus the most prominent feature is the black 
keel-like posterior ridge, while the anterior portion mainly consists 
of the thickening (/. a .). 



Fig. 6 . Gloss ina fuscipleuris : $. Signura, X c.ioo. A. flattened specimen, dorso- 
ventral aspect; B. folded condition ; C. flexed condition, lateral aspect; /., lamina ; l.e ., 
lateral expansions of posterior projection ; t.a ., anterior thickening; t.p., posterior thickening. 


Type of female armature from the Belgian Congo (ex-coli. 
Mtisee Royal d’Histoire Naturelle de Belgique); no further data 
available. Two others were taken at Daru Forest, Uganda 
Protectorate, 25-29.10.1911 (Dr. S. A. Neave), and one at Buamba 




4 * 


Forest, Semliki Valley (2,300-2,800 feet), 3-7.11.19n (Dr. S. A. 
Neave). In the latter specimen the signum was doubled on itself 
(fig. 6, B), and the posterior pair of thickenings (/. p.) were much 
shorter and less prominent than in the other specimens examined. 


Glossina tabaniformis , Westwood 

External armature of the female (fig. 7). The dorsal plates in 
this species resemble those of G. fusca, but the greatest width may 
be equal to one-third the length instead of one-quarter, which is the 
average ratio in G. fusca. The outer margin is strongly angled at 
the point of greatest width. The anal plates are rotated forwards 



to a marked degree so that their free edges project ventrally, and 
when mounted with slight pressure they obscure from view the sternal 
plate. The condition has also been observed in isolated specimens 
of other species, but it is only in G. tabaniformis that it appears to be 
a constant character. 

Signum of the uterus (fig. 8). The signum consists of a lyriform 



43 


lamina somewhat depressed medianly. A transverse constriction 
divides it into two unequal portions, the proximal roughly twice as 
long as, and one and a half times as broad as, the distal. The 
former is sub-ovate in outline and bears two or three terminal 
processes diverging to a varying extent. The anterior division 
terminates in two projections. The whole is strengthened by a pair 
of curved bands of thickened chitin. 



Fig. 8. Glossina tabanijormis : $. Signum, X e. 155. 


Type of female armature taken at Yombi Yombi, Leverville, 
Congo, 3.10.13 (Dr. Arbrassart). Two were taken at Yakusu (?), 
Haut Congo (Rev. Sutton-Smith) and another in Nigeria (Dr. T. R. 
Leonard). : _ ... : 





44 


Glossina s ever ini , Newstead 

External armature of the female (fig. 9). The characteristic 
feature of the armature of this species is the great extent of the dorsal 
plates. The external borders are strongly convex; the internal 
distally straight, and closely approximated in the median line, for 
a distance of half the total height of the armature. Proximally the 
latter diverge to form the concave inner edges of the ventral limb (y.) 
of the plate. The other plates are similar to those in G. fusca. 



Fio. 9. Glossini scverini: External armature, X c. 80. v. } ventral limb of dorsal plate. 


Signum of the uterus (fig. 10). The outline of the signum in 
dorso-ventral aspect (fig. 10) is oblong, the length being one and a 
half times the greatest width. It is composed of a pair of plates 
separated throughout their length . Each plate consists of an 
elongate proximal limb surmounted by a broader distal piece, the 
latter medianly closely approaching that of the opposite side. 






45 

Outwardly the plates are curved upwards so as to embrace the lateral 
walls of the genital fossae. 

Type of female armature from the Belgian Congo (ex-coll. Mus£e 
Royal d’Histoire Naturelle de Belgique); no further data available. 



Fig. io. Glossina severtni: $. Signum, x c, 155. 


In a second specimen from this region the ventral limb of the dorsal 
plate was more prolonged than in that of the one figured here. 




Glossina medicorum , Austen 


External armature of the female (fig. il). The dorsal plates are 
of a type frequently met with in G. fusca. They are almost 
uniformly narrow, the long axes slightly convex, and the extremities 
obliquely truncate. The lateral plates are inclined towards one 
another, and when mounted largely obscure from view the ventral 
plate. 



Fio. n. Glossina me dicorum : $. External armature, x r.8o. 


Signum of the uterus (fig. 12). The outline of the fossae in 
dorso-ventral aspect is elongated; the proximal portion cordiform, 
an appearance produced by the presence of two convergent ridges, 
separated by a deep median cleft. The length is twice the greatest 
width. The signum consists of two distally convergent ochraceous 
bands which clothe the crests of the two ridges and posteriorly are 
sharply curved to meet one another in the median line. A pair of 
crenulated, chitinous thickenings ( c . r.) of the wall of the fossae 
extend from the narrow anterior extremities of the sclerites toward 
the border of the fossae, and in the region of their termination lie one 
or two irregular groups of dark granules. All four specimens 
exhibited to a well marked degree the aristal character described by 
Newstead (1913). 





Type of female armature, from Black Volta River, N. Ashanti, 
13.4.10 (Dr. A. Kinghom). A second was without data; a third 
was from Volta River, P.A. Gl. Rapids, G.F.S., 1.12 (Dr. A. M. 
Dowdall), and a fourth from Volta River, 30. m. N. Kpong, 9.1911, 
Gold Coast (Dr. P. D. Oakley). 


Fig. 12. Glouina mediocrum : $. Signum, X c. 155. cr., crenulated thickening of wall 
of genital fossa. 






4 8 

Glossina brevipalpis t Newstead 


External armature of the female (fig. 13). The dorsal plates are 
relatively broader than in any other members of the group, the 
greatest width being about two-thirds the greatest length. The 
obliquely-placed inner edges are almost straight. The margin is 
curved externally, bluntly pointed apically and obliquely truncate 
proximally. Spines are confined to the inner halves of the plates. 
The lateral plates are broad and their free edges approach one 
another medially. In the membrane at the base of the ventral plate 
occurs a pair of hamate sclerites (A. s.\ the outer lateral edges of 
which are irregularly dentate. 

Signum of .the uterus . Absent. 



Fig. 13. Glossina brevipalpis: External armature, x r.8o. fu hamate tderite. 


The type of female armature was taken at Ng’ani Nyassa, 1912 
(Dr. : M! Sanderson); two examples at Katanga* Belgian Congo 
(Dr. J. Schwetz), another Juba River, East Africa (Dr. R.'P*. Filleul) 
and a fifth at Makulu, Congo, 11.7.04 (Drs. Dutton and Todd). 




+9 

Glossina longipennis , Corti 

External armature of the female (fig. 14). The external armature 
is unusually prominent, owing to the fact that the dorsal plates do 
not lie in the ’ same plane, but are opposed to one another at 
a considerable angle, carrying out the lateral plates, which project 
from the extremity of the abdomen to a very marked extent. The 
dorsal plates are sub-triangular in outline, and their inner edges are 
adjacent from the apex to a distance approaching one-third of the 
length of the plate. Setae are confined to the proximal two-thirds of 
the inner half of each plate. 

Signum of the uterus Absent. 



Type of female armature from Nairobi, British East Africa, 
28.5.12 (Dr. A. D. Milne). In two other examples examined, one 
from this region, a second from Entebbe, Uganda (Dr. Christy), the 
dorsal plates were entirely membraneous; their position merely 
indicated by the presence of the dark spines. 



5 <> 


GROUP II. PALPALIS Group (figs. 15, 16). External armature , 
consisting of six plates : in addition to those present in 
Group /, there is a small medio-dorsal plate. Signum of 
uterus absent . 

External armature . Mean dimensions o*6 x 0*5 mm. This 
structure presents an essentially compact aspect; the various plates, 
which are of comparatively great superficial extent, are of such form 
and disposition that, together they almost entirely fill the space in 
which the armature lies. The dorsal plates (< d . p.) are sub-triangular 
in outline, their inner edges lying parallel and in close proximity. 
They bear spines smaller but similar in character to those borne by 
the dorsal plates of G . fusca. The anal plates (a. p.) are broadly 
triangular, and are closely applied to the underlying integument, to 
which they are attached by all but their free inner edges. Thus they 
cannot be reflexed as can those of the Fusca Group. Between the 
lateral and dorsal plates occurs a small oval sclerite ( m.d.p.\ which 
possibly corresponds to the 1 dorsal scale ’ of the Blow-fly, but which 
is absent in the Fusca Group. It may, on the other hand, be a 
detached median portion of the VIII° tergite. 


Glossina palpalis , Robineau-Desvoidy 

External armature (fig. 15). The outline of the containing area 
is roughly circular. The dorsal plates ( d . p.) are of chitin varying 
from brown to pale ochraceous colour, in one case they were as 
transparent as the surrounding membraneous integument. The general 
shape is that of a quadrant, the curved border lying externally, 
and the vertical inner edges parallel and closely approximated. 
The latter are sometimes almost entire, but frequently a variable 
number of indentations causes a jagged and irregular outline. In 
some cases the two plates may be connected medially by one or more 
bridges of chitin. The transverse proximal border of each dorsal 
plate is usually to a greater or less extent emarginate in the middle 
of its length, with the result that the inner angular portion, which is 
devoid of long spines, is more or less constricted off from the rest of 
the plate. The anal plates (a. p.), when mounted, consist of a 
broadly sub-triangular portion which lies almost parallel with the 



integument, and is folded externally so as to overlie the smaller basal 
portion from which it arises. The free edge is directed towards the 
sternal plate and the distal border appears to have become 
secondarily fused with the integument. The sternal plate ( st . p.) is 
broadly rounded apically and, as in the lateral plates, the basal 
portion lies below that seen in surface view. The width at the base 
is greater than the length. The medio-dorsal plate is variable in 
size, and may consist of one, two, or three sclerites. 



Fig. 15. Glossina palpalis : $. External armature, X r.90. m.d.p., medio-dorsal plate. 


Type of female armature taken at Kintampo, N. Ashanti, April 
to July, 1913 (Dr. A. Ingram); ten other specimens from this 
locality were examined. Eight were taken at Volta River, Gold 
Coast, 1913 (Dr. A. M. Dowdall), and two of a ‘small dark grey 
form', Illorin, N. Nigeria (Dr. J. W. Scott Macfie). 


Glossina palpalis , Rob.-Desv., race fuscipes , Newst. 

External armature of the female. Resembled that of G. palpalis , 
but the constriction of the angular portion of the dorsal plate was 
very well marked. Further, the presence of well developed spines in 
this median region was a constant feature, and frequently these were 
arranged in a definite ‘ V * shaped group, the apex directed dorsally 
and lying in the median line between the dorsal plates at about the 



5 * 


middle length. The presence or absence of these medianly-placed 
spines can usually be determined by examination of the terminal wall 
of the untreated dry specimen under the low power of the microscope. 
Before attributing any definite value to this character, however, it is 
hoped by examination of further material to discover whether or not 
this course is justifiable. 

Six specimens examined were taken at Nimule, Uganda, July, 
1911 (Dr. R. G. McConnel). 


Glossina caliginea t Austen 

External armature of the female . Examination of four specimens 
revealed no features of distinction which could be used to separate 
this species from G. palpalis . The examples were taken at Yewa 
River, South Nigeria, 10-12.9.1911 (Captain L. E. H. Holmfrey). 


Glossina pallicera y Bigot 

External armature of the female. The dorsal plates, unlike 
those of G . palpalis , are not right angled internally, but in one 
specimen the two shorter sides of the plate enclose a widely 
obtuse angle, and thus expose above the anal and sternal plates 
a comparatively large triangular median area of unchitinised 
membrane. In the other specimen examined the dorsal plates are 
bounded by a continuous curved margin, the outline of the plate 
being ovoid. In both cases the greatest width of the plate was 
one-half of the greatest length, while in G. palpalis this ratio is 
approximately 2:3. The sternal plate was, in both examples, 
proportionally longer and narrower than that of any other member 
of the Palpalis Group, the width at the base being considerably less 
than the greatest length; in the second specimen it was markedly 
attenuated and digitiform in outline. The medio-dorsal plate was 
of very small extent in both cases. The first example was taken at 
Cote dTvoire, 1910 (Prof. E. Roubaud, ex-coll. Museum, Paris), the 
second at Tetchari, Tanoso, W. Ashanti, 4.8.10 (Dr. A. Kinghom). 



53 

Glossina tachinoides , Westwood 

External armature of the female (fig. 16). Dorsal plates right 
angled internally but much narrower and more elongated than in 
G. palpalis . The height is rather less than twice the greatest width. 
The other plates resemble those of G. palpalis. 



St. p — 


Fig. 16. Glossina tachinoides : $. External armature, X c. 90. 


Type of female armature taken at Lorha, Gold Coast, 1915 
(Dr. J. F. Corson); a second specimen bore the same data. Other 
specimens were from Salago, Gold Coast, June 12th (Dr. Lefanu). 


GROUP III. Morsitans Group (figs. 17, 18). External armature , 
consisting of a pair of fused anal plates and a median sternal 
plate . Dorsal plates generally absent. Signum of uterus 

absent . 

External armature . Mean dimensions 0*4 x o*2 mm. The 
condition and extent of the armature exhibit very considerable 
reduction from that of the Fusca Group. In consequence of the loss 
or reduction of the dorsal plates, the area in which the armature lies 
is much shallower than in the case of Groups I or II, and is roughly 



54 


in the form of an ellipse, the long axis of which runs transversely. 
The anal plates (a.p.) are of almost membraneous character, the walls 
collapsing when mounted under slight pressure after treatment in 
boiling KOH. They are united dorsally to form a continuous arc 
surrounding the free edge of the sternal plate. The median portion 




Fig. 17. Clossina morsitans : $. External armature, X c. 155. A. mounted without 
maceration j B. mounted with slight pressure after maceration. 


of the arc (;//. d. p.) projects dorsally as shown in fig. 17, B, and 
possibly represents the 4 medio-dorsal scale*, which has become fused 
laterally with the anal plates. G. austeni was exceptional in the 
possession of a pair of small, though well developed dorsal plates. 




55 


Glossina morsitans, Westwood 

External armature of the female (fig. 17). Dorsal plates generally 
absent, but when present extremely thin, not bearing dark spines. 
For other characters see pp. 53-4. 

Nine of the examples were taken by Prof. R. Newstead and 
Dr. J. B. Davey during the Nyasaland Expedition of 1911. Seven 
others were from Zambesi, 1907 (Kinghom and Montgomery). 

Glossina austeni, Newstead 

External armature of the female (fig. 18). This is distinguished 
from that of all other members of the Morsitans Group by the 
possession of a pair of small, but well defined, dorsal plates ( d.p . and a) 
bearing relatively very long spines, the latter of a similar character 




Fig. 18. Glossina austeni: External armature, x r.155. A. dorsal plate, X f.240. 


to those borne on the dorsal plate in G. fusca. The dorsal plates 
are sub-triangular in form and widely separated. 

The three examples examined were taken at Jubaland, 20.8.12 
(Dr. R. P. Filleul). 



56 


Other species of this group were investigated, but none of these 
shewed any marked features distinguishing the armature from that 
of G. morsitans. The localities from which they were taken were as 
follows: — 

G. submorsitans, Newstead. Makongo, Gold Coast, 14.10.14 
(Dr. J. F. Corson); Prang, Gold Coast, 10.10.14 (Dr. J. F. Corson). 

G. pallidipes, Austen. Two specimens at Juba River, East 
Africa (Dr. R. P. Filleul), and two others at Alexandra, Gosha, 
Jubaland, 6.1912 (Dr. R. P. Filleul). 

G. longipalpis, Wiedemann. Two specimens taken at Edinam, 
Ashanti, 3.5.13 (Dr. F. H. StoreyV 


REFERENCES 


Cowne, B. T. (1893-95). The Blow Fly, Vol. II, pp. 745-6 and 675-77. 

N’f.wstead, R. (1911). A Revision of the Tsetse-flies ( Glossina ), based on a Study of the Male 
Genital Armature. Bull. Ent. Res., Vol. II, May, 1911. 

- (1913). A New Tsetse-fly from the Congo Free State, and the Occurrence of Glossina 

austeni in German East Africa. Ann. Trop. Med. & Parasitol ., Vol. VII, No. 2, June, 
1913. 

Pierce, F. N. (1914). The Genitalia of the Geometridae 

Stuhlm4 NN, F. (1907). Beitrage rur Kenntniss drr Tsetsefliege Arbeiten aus. dem. Kaiserltcben 
Gesund , Bd. XXVI> Heft. 3, pp. 57, 58. 



57 


STRONGYLIDAE IN HORSES 

VII. CYUCOSTOMUM PATBRATUM sp. n. 

BY 

WARRINGTON YORKE 

AND 

J. W. S. MACFIE 

(.Received for publication io March , 1919) 

Size and Shape. A moderately small species of the Genus 
Cylicostomum , the female being slightly larger than the male. 
Twenty females and ten males were measured. The males were 
from 8 0 to 9 5 mm. in length, average 8*6 mm.; the females from 
8*4 to 11 mm. in length, average 9 5 mm.; the greatest breadth, in 
those worms which were properly orientated, averaged, males 380/1, 
females 393 /1. 

Head. A well-marked neck separated the head from the body. 

Mouth collar. Marked off from the rest of the skin by a definite 
constriction. 

Head papillae. Sub-median, pointed and projecting, their 
extremities separated off from the remainder by lateral notches; 
lateral, prominent. 

Mouth capsule. Ellipsoidal in transverse section, the ratio of 
the lateral diameter to the dorso-ventral diameter of the anterior 
opening of the buccal capsule being about 1 to 1*3. When the 
worm is properly orientated the walls of the mouth capsule seen 
in optical section are wedge-shaped, being slender anteriorly and 
stout posteriorly; the inner surface is deeply notched at the level 
of the insertion of the internal leaf crown (fig. 1). When viewed 
laterally the walls of the buccal capsule converge considerably from 
before backwards (fig. 3). In properly orientated worms the 
antero-posterior diameter (i.e., the distance from the anterior to 


I 



5 « 

the posterior opening) of the buccal capsule varies in the males from 
25 fi to 28*5^, average 267ft, and in the females from 26*5ft to 28/1, 
average 27*2/4. When the worm is lying on its side the mouth 
capsule appears less deep owing to the fact that the walls of the 
buccal capsule are set obliquely (fig. 3). In the males the lateral 
diameter of the buccal capsule at the anterior opening varies from 



Fig. 2 



Figs. 1-2. Cylicostmnum falrratum sp. n. 
Anterior extremity, ventral view. Fig. i X 360. Fig. 2 X 90. 


8o/i to 8 7/1, average 85 /i, and at the posterior opening from 75/* to 
84 ft, average 79 ft; in the females the lateral diameter of the buccal 
capsule at the anterior opening varies from 78/1 to 97 /i, average 89/4, 
and at the posterior opening from 8o/i to 95 /u, average 88/4. The 
ratio of the lateral diameter of the anterior opening of the buccal 




59 


capsule to that of the posterior opening is therefore about i to I. 
The ratio of the lateral diameter of the buccal capsule at the anterior 
opening to the antero-posterior diameter is in both sexes about 
3 25 to 1. 



Fic. 3. Cylicostomum fateratum sp. n. 
Anterior extremity, lateral view, x 360. 


Dorsal oesophageal gutter . Does not project into the buccal 
capsule. 

Leaf crowns . The external leaf crown consists of about twenty- 
four large pointed elements arising from the mouth collar. The 



6o 


internal leaf crown consists of numerous long narrow elements 
arising from a sinuous line situated deep in the mouth capsule 
(figs, i and 3). 

Oesophagus. The length in ten males varied from 530/1 to 602/1, 
average 559/1, and the greatest breadth from 131/1 to 155/“, average 
146/1; the ratio of breadth to length is 1 to 3 8. In twelve females 
the length ranged from, 561/1 to 595/1, average 582/1, and the greatest 
breadth from 136/1 to 171 /*, average 149/1; the ratio of breadth to 
length is 1 to 3 9. The ratio of the length of the oesophagus to that 
of the worm is, in the male 1 to 15, and in the female 1 to 16. 



Figs. 4-8. Cylicostomum pateratum sp. n. 

Fig. 4: Posterior extremity of male, ventral view, X 90. 

Fig. 5 : Posterior extremity of male, lateral view, X 90. 

Fig. 6: Genital appendages, ventral view, X 360. 

Fig. 7: Genital cone and appendages, lateral view, X 360. 

Fig. 8 : End of spicules X 1360. 

Excretory Bladder. Lies just behind the nerve ring. The 
distance of its posterior margin from the posterior end of the 
oesophagus varies from 208/1 to 280/1, average 244A 1 * 

CERVICAL Papillae. Lie at about the same level as the excretory 
bladder. 



6i 


Posterior Extremity of Male. The dorsal lobe of the bursa is 
short, about semicircular (fig. 4). In eight worms the length of the 
main trunk of the posterior ray from the tip to the point of origin of 
the postero-external rays varied from 391 /1 to 495/1, average 442/1. 
The ratio of the average length of the main trunk of the posterior 
ray to the average length of the male worm is 1 to 19*5. 

Genital Cone. The dermal collar is well developed on both the 
ventral and dorsal surfaces of the genital cone. The genital 
appendages on each side are represented by slight elevations 
furnished with three conical processes, the innermost bearing a long 
delicate finger-like process (fig. 6). 

Spicules. The ends of the spicules are barbed as shown in fig. 8. 



Fig. 9. Cylicostomum pateratum sp. n. 
Posterior extremity of female, lateral view, x 90. 


Posterior Extremity of the Female. The end of the body is 
bent dorsally at right angles. The ventral prominence is large and 
projecting. The tail is very short and conical (fig. 9). In six 
worms the distance between the anus and vulva varied from 91 At to 
116/1, average 104/1, and the distance measured straight along the 
middle line of the tail from the tip to a line drawn horizontally 
through the anus varied from 105/1 to 143 /*, average 114/1. 



62 


DIAGNOSIS. The following are the chief diagnostic characters of 
this worm: — 

1. Size, moderately small; average length, males 8'6 mm. and 
females 9'5 mm. 

2. Buccal capsule: anterior opening ellipsoidal, ratio of lateral 
to dorso-ventral diameter of anterior opening of capsule is 1 to 1*3; 
walls when seen in optical section, in properly orientated worms, are 
wedge-shaped, being slender anteriorly and stout posteriorly; the 
inner surface is deeply notched; ratio of breadth at anterior opening 
to antero-posterior diameter 3'25 to 1. 

3. Dorsal oesophagal gutter does not project into buccal 
capsule. 

4. Dorsal lobe of bursa short, almost semicircular; ratio of 
length of posterior ray to total length of male worm I to 19' 5. The 
genital appendages are slight elevations furnished with three conical 
processes, of which the innermost is largest and bears a long delicate 
finger-like process. 

5. Termination of female body bent dorsally at right angles; 
ventral prominence large and projecting, tail short and conical. 

This species clearly belongs to the catinatum-alveatum group. 
In the character of the mouth capsule it most closely resembles 
C. goldi, but can be distinguished from this worm by its larger size, 
by the appearance of the walls of the mouth capsule when seen in 
optical section, by the fact that the line of origin of the internal leaf 
crown is sinuous and situated at a much deeper level than in 
C. goldi, by the fact that the tail of the female is bent dorsally at 
right angles whereas in C. goldi this is not quite the case, and by 
the appearance of the accessory bodies of the genital cone. 



63 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXII. INTRAMUSCULAR INJECTIONS OF 
QUININE BIHYDROCHLORIDE GRAINS 15 ON 
EACH OF TWO CONSECUTIVE DAYS ONLY, 
IN MALIGNANT TERTIAN MALARIA 

BY 

Lif.ut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 
C. FORSTER COOPER 

AND 

H. F. CARTER 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Oifice 
(Received for publication 2 February, 1919) 

In previous studies (1917 and 1919) we have recorded the results 
of intramuscular injections of quinine bihydrochloride in simple 
tertian malaria. The present observations refer to malignant tertian 
malaria. 

The treatment was an intramuscular injection of quinine 
bihydrochloride (grains 15 in 2 c.c. of water) on each of two 
consecutive days only. All the cases were adult* males infected 
either in West Africa (Dakar) or in Macedonia. 

In every instance a diagnosis of malignant tertian malaria was 
made microscopically, and in all cases trophozoites were present in 



6 + 


Case 1353 


|*0 |2» 12? 1 23 I 24 [aS |2fe | 27 12> \lq [30 | 


-«I" «i* 


ci 0lJ 


Cl Itct-I O I Cl I t Itc« I 


CAiE I355 












6S 


Case 1359 


D*c- 



Case 1363 


Jaw 














66 


Tam* 


Summary o! results of intramuscular injections of tjuinine grains 15 on each of two consecutive days only in malignant tertian mdca 

• S. « Salonika. W.A. — West Africa. 


Number 

of 

case | 

: 

•Place 
of in¬ 
fection 

Interval 
[in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after first 
injection 

Trophoxoites 

disappeared 

from 

cutaneous 
blood 
in — days 
after first 
injection 

Trophozoite 
relapse 
occurred 
in — days 
after first 
injection 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after first 
injection 

Remark. 

1347 

S. 

... 

... 

... 

*2.3.17 

Apyrexia 

2 

8 

No record 

Crescents 00 8th r 

1348 

S. 

... 



16.3.17 

3 

1 

18 

... 

Crescents penury: 

tidily thrauh- 
Quinine orally. ess 
on 20th day 

1349 

S. 



... 

3 - 4*7 

Apyrexia 

1 

*7 

18 


1350 

W.A. 

1 

1 

0 

17.11.17 

Same day 

2 

18 

>9 

Crescents 00411-:: 
I3th-I5th 2 s.:is 

1 35 1 

W.A. 

1 

1 

0 

17.11.17 

1 

4 

>3 

18 

Crescents on 511^1 
and 12th dirt 

* 35 * 

W.A. 

1 

1 

0 

17.11.17 

1 

3 

8 

11 

Crescents on 6ti z 
day*. 

*353 

W.A. 

1 

1 

0 

22.11.17 

3 

2 

5 

8 

Crescents cm is: r 
4th-7th days. 

>354 

W.A. 


1 

0 

22.11.17 

2 

2 

8 

11 

Crescents on z=c z 
6th days. 

>355 

W.A. 

1 

1 

0 

23.11.17 

1 

2 

17 

18 

Crescents on r:: 
9th-uth dip. 

1356 

W.A. 

t 

1 

0 

28.11.17 

2 

3 

>7 

16 

Crescents on ni: 
6th-9th, nti J 

16th days. 

>357 

W.A. 

1 

1 

0 

: 

28.11.17 

Apyrexia 

2 

6 

9 

Crescents on aa-r 

1358 

W.A. 

1 

1 

0 

6.12.17 

Same day 

2 

21 

22 

Crescents on 6c. 3 
and nth-2is^' 

>359 

W.A. 

1 

1 

0 

7***7 

2 

2 

13 1 

*4 


1360 



... 

... 

17.12.17 

1 

2 

7 

1 9 

Crescents on 211- 
and 7th dm 

1361 

... 


... 

... 

2.1.18 

Apyrexia 

2 

>5 

>5 

Crescents onisi- 
6th-12 th ini u= 

1362 

... 


1 

... 

2.1.18 

Apyrexia 

1 

6 

>5 


>363 

... 

... 

i ... 


2.1.18 

1 

1 

3 

*3 

Crescents on is i- 

1364 

S. 

37 

: • 

1 

28.8.18 

2 

2 

>5 

1 >6 


>365 

S. 

26 

1 

0 

23 - 9 «* 

1 

2 

7 


Crescents on i»i-~ 

1366 

W.A. 

1 

1 

0 

23 . II. l8 

2 

2 

>4 

>6 


1367 

W.A. 

1 

1 

0 

23.II.l8 

2 

2 

, >5 

>9 

Crescents on xsc- 

1368 

W.A. 

1 


0 

25 . II. l8 

2 

2 

20 

*3 

Crescents on 4th- ■ 
13th-18th dap 

>369 

S. 

5 

l 3 

1 

1 2.12.18 

2 

2 

9 

10 

Crescents on xai 1 

>370 

W.A. 

2 

2 

1 

2.12.18 

1 

2 

>7 

18 

1 

1371 

S. 

4 

3 

2 

! 3.12.18 

i 

Apyrexia 

1 

7 

i 

Crescents on pi ft 
Quinine orally. p 
on 8th day. 

> 37 * 

s. 

1 

1 34 

3 

2 

' 3.12.18 

Apyrexia 

1 

16 

1 

Quinine oraDp P 
on 17th day. 

>373 

W.A. 

1 1 

1 

| 

0 

! 3.12.18 

| > 

i 

1 

>9 

20 

Crescents on rsd * 
18th and 19th P 

>374 

s. 

16 

1 

1 

1 

4.12.18 

Apyrexia 

! 2 

7 

8 

Crescents 00 ph - 

>375 

s. 

3 

1 

1 4 

3 

1 4.12.18 

1 

Apyrexia 

| 1 

8 

1 

... 

Quinine orafir. 
on 10th day. _ 





67 

the blood on the day treatment commenced. Blood examinations 
were made daily. 

In the temperature charts: — 

t. = malignant tertian trophozoites. 

cr. = crescents. 

neg. = no parasites found 

Q.M. = intramuscular injection of quinine bihydrochloride grains 15. 

In nine of the twenty-nine cases (Nos. 1347-1375) treatment 
commenced during an apyrexial period; in the remaining twenty 
cases the temperature fell to normal within three days. Trophozoites 
disappeared from the cutaneous blood within four days. Observa¬ 
tions regarding the crescents will be found in the ‘ Remarks * column 
of the table. 

Relapses 

A parasitic relapse occurred in all cases, trophozoites reappearing 
in the cutaneous blood in three to twenty-one days; in Cases 1353 
and 1363 trophozoites were absent for respectively three and two 
days only. Febrile relapses occurred in eight to twenty-three days. 


CONCLUSION 

An intramuscular injection of quinine bihydrochloride grains 15 
in 2 c.c. of water on each of two consecutive days only, causes the 
cessation of febrile paroxysms and effects the disappearance of 
trophozoites from the cutaneous blood in malignant tertian malaria. 
The action, however, is only temporary, a relapse occurring within 
three weeks, occasionally within a few days. 


REFERENCE 


Stephens, J. W. W., Yorke, W., Blacklock. B., Macfie, J. W. S., Cooper, C. F., and 
Carter, H. F. (1917 and 1919). Ann . Trop. Med. & Parasitol. , Vol. XI, p. 113, and 
Vol. XII, p. 402. 




69 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXIII. ORAL ADMINISTRATION OF QUININE 
SULPHATE GRAINS 30 ON EACH OF TWO 
CONSECUTIVE DAYS WEEKLY, OVER A 
PERIOD OF FIVE WEEKS, IN MALIGNANT 
TERTIAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 
C. FORSTER COOPER 

AND 

H. F. CARTER 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
(Received for publication 9 February , 1919) 

All the cases were adult males, infected in West Africa. In 
every instance a diagnosis of malignant tertian malaria was made 
microscopically, and in all cases trophozoites were present in the 
blood on the day treatment commenced. Blood examinations of 
ordinary thin films were made daily. 

The records of the observations are given in the tables, in 
which: — 

o = absence of fever and trophozoites. 

1, 2, etc. = number of trophozoite febrile relapses weekly. 

I*, 2 # , etc. = number of non-trophozoite febrile attacks weekly. 
t 1 , t\ etc. = number of non-febrile trophozoite relapses weekly. 
c\ c*, etc. = number of days on which crescents were present weekly. 

Note.— A rise of temperature above ioo° F., of which the nature is 
unknown is termed a febrile attack . A similar rise of temperature accompanied 
by trophozoites in the blood at the time, or within three days, is termed a 
trophozoite febrile relapse or true relapse . The term paroxysm is used indifferently 
to denote any febrile disturbance of ioo° F. or more. 



Table I. 


Results of oral administration of quinine sulphate grains 30 on each of two consecutive days 
weekly for 5 weeks, in malignant tertian malaria. 


1 

i 

- 1 

Number 

of 

case 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
irt — days 
after 

Trophozoites 

disappeared 

from 

cutaneous 
blood 
in — days 

Number of febrile paroxysms (trophozoite 
and non-trophozoite) and results of blood 
examinations during treatment 

Week of treatment 

1 

' 

first dose 

after 
first dose 

I St 

2nd 

3 rd 

4th 

5th 

1 

1376 

24.11.17 

1 

3 

ole* 

0 

cl 

O 

cl 

2 

t 3 C * 

>377 ' 

29.11.17 

1 

2 

0 

0 

cl 

O 


1* 

O 

> 37 * 

29.11.17 

3 

2 

0 

0 


O 


0 

O 

*379 

29.11.17 

1 

4 

0 

0 


O 


0 

O 

1380 

29.11.17 

2 

2 

0 

0 


O 


0 

O 

1381 

29.11.17 

Same day 

4 

0 c* 

0 

c* 

O 


O cl 

O 

1382 

29.11.17 

3 

2 

0 c 1 

0 


0 


0 

O 

1383 

29.11.17 

Same day 

i 2 

O C l 

0 


0 


0 

O 

1384 

1.12.17 

« 

2 

0 

0 


0 


0 

O 

1385 

1.12.17 

1 

2 

0 


I* 

0 


0 

O 

1386 

6.12.17 

1 

2 

0 c* 

0 


0 


! 

O 

O 

1387 

, 3- I2 - I 7 

1 

2 

O C* 

0 

c» 

0 

cl 

0 ci 

O 

1388 

13.12.x7 

Apyrexia 

2 

0 c- 


I* 

tl 


0 ! 

1 

O 

>389 

13.12.17 

1 

2 

0 c* 

0 

cl 

0 


0 

O 

> 39 ° 

1 3* I2 - I 7 

Same day 

3 

0 

0 


0 


0 

O 

* 39 * 

20.12.17 

1 

r 

0 

0 


0 


0 

O 

> 39 * 

17.11.17 

2 

2 

0 

0 

C* 

0 

c* 

... 

... 

>393 

29.11.17 

2 

4 

0 cl 

0 

d 

0 

cl 


... 


t Four days are allowed for the initial fever to subside and for the disappearance of 
trophozoites from the blood. The absence of fever and trophozoites in this column refer 
to the last two days of the week only. 




71 


Table II. 


Summary of Table I. 


Week of treatment 

ISt 

2nd 

3rd 

1 

4th 

5 th 

NumlMr of ohm treated . 

EM 

It 

18 

It 

16 

Number of cate* having trophozoite febrile relapses 


o 

1 ° 

HI 

o 

Number of cases having non-trophozoite febrile 
attacks . 


2 1 

! 

1 o 

. 

■ 

o 

Grand total of all fobrlie oases . 


2 

0 

2 

0 

Total number of trophozoite febrile relapses 

o 

o 

o 

2 

o 

Total number of non-trophozoite febrile attacks 

o 

2 

o 

X 

o 

Grand toad of all febrile paroxysms 

0 

2 

0 

3 

0 

Number of trophozoite cases(febrile and non-febrile) 

o 

o | 

1 

i 

I 

Number of crescent cases . 

9 

t 

7 

4 

2 

1 


Table III. 


Analysis of Table II. 


Week of treatment 

! 

> ISt 

1 

2nd 

3 rd 

4 th 

5th 

Average 

per 

week 

Percentage of trophozoite febrile 
relapse cases per cases treated' 

0 

o 

o 

6o 

o 

It 

Percentage of all febrile (trophozoite 
and non-trophozoite) cases per cases 
treated . 

o 

IPO 

■ 


° 1 

47 

Percentage of all trophozoite (febrile 
and non-febrile) cases per cases 
treated . 

o 

I 

o 


H 

6*0 

3 * 

Percentage of crescent cases per cases 
treated .j 

D 

390 

22*0 

12-5 

6*o 

26*0 



























72 


In one of the eighteen cases (Nos. 1376-1393) treatment was 
commenced during an apyrexial period; in the remaining seventeen 
cases the temperature fell to normal either on the same day or in one 
to three days. 

Trophozoites disappeared-from the blood in one to four days. 
Relapses 

During treatment . Owing to the discharge from hospital of two 
cases in the fourth week, the number of cases treated was eighteen in 
the first four weeks and sixteen in the fifth week. 

The average weekly number, over a period of five weeks, of cases 
which had (1) trophozoite febrile relapses was V2 per cent, of cases 
treated, (2) non-trophozoite febrile attacks 3*5 per cent., (3) febrile 
paroxysms, trophozoite and non-trophozoite, 4*7 per cent., (4) tropho¬ 
zoite relapses, febrile and non-febrile, 3*5 per cent., and (5) crescent 
relapses; 26 per cent: 

After treatment. The cases were not observed. 


SUMMARY 

As a palliative, quinine sulphate grains 30 on each of two 
consecutive days weekly, over a period of five weeks, suffices to keep 
the blood free from trophozoites and to prevent relapses in the great 
majority of cases. It is noteworthy that the percentage of cases 
having crescents in the peripheral blood diminishes each week, viz., 
frpm 50 per cent, in the first week to 6 per cent, in the fifth week of 
treatment. 



73 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXIV. THE DISAPPEARANCE OF CRESCENTS 
UNDER QUININE TREATMENT 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 
C. FORSTER COOPER 

AND 

H. F. CARTER 

Front the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
{Received for publication 14 February , 1919) 

In a previous study (1917) we have referred to various 
contradictory statements in the literature on this subject. We record 
here observations made by us on eighty-nine crescent cases.. 

The cases were divided into three groups according to the daily 
dose of quinine administered : — 

Group A. Quinine sulphate in solution orally, grains 20 daily. 

(Cases 1394-1413). 

Group B. Quinine sulphate in solution orally, grains 30 daily. 

(Cases 1414-1451). 

Group C. Quinine sulphate in solution orally, grains 45 daily. 

(Cases 1452-1482). 

All the cases had crescents in the blood on the day treatment 
commenced. Blood examinations were made once weekly, thick 
films being used. As these observations were made in 1916 at the 
beginning of our work on malaria, information as to place and 
duration of infection is not available. 

The results are given in the table, from which it will be seen that 
at the end of four weeks* treatment crescents had disappeared from 
50 per cent, of cases treated when the daily dose of quinine was 
grains 20, and from over 90 per cent, when the daily dose was 
grains 30 or 45. 




74 


Tabls 

Showing the disappearance of crescents under quinine treatment. 
A. Quinine sulphate grains 20 daily. (Cites 1394-1413). 



At 

beginning 

Week of treatment 


of 

treatment 

1st 

2nd 

3rd 

4 th 

Number of cases observed . 

20 

20 

20 

20 

20 

• 

Number of cases showing crescents . 

20 

*9 

18 

*3 

j ,0 

Percentage of cases showing crescents. 

IOO% 

95% 

90% 

63% 

50% 

B. Quinine sulphate grains 30 daily. 1 

(Caiet 1414-1451) 




At 

beginning 

Week of treatment 


of 

treatment 

ISt 

2nd 

3 rd 

4th 

Number of cases observed . 

3* 

38 

38 

38 

38 

Number of cates showing crescents . 

3* 

2 9 

*7 

8 

3 

Percentage of cases showing crescents. 


76% 

45 % 

*« % 

** 

00 


C. Quinine sulphate grains 45 daily. (Cases 1452-1482). 



At 

beginning 

Week of treatment 


of 

treatment 

1st 


3rd 

4 th 

Number of cases observed . 

31 

H 

3 * 

3 * 

3 * 

Number of cases showing crescents . 

3 1 

20 

*4 

11 

2 

1 

Percentage of cases showing crescents . 

100% 

65 % 

45% 

1 35% 

6 % 


CONCLUSION 

Under quinine treatment grains 30 or 45 daily, crescents do not 
persist in the cutaneous blood in the majority of cases for more than 
three weeks. 

REFERENCE 

Stephen**, J. W. W., Yorkf., \Y., Bucklock, B., Macfif, J. YV. S., and Cooper, C. F. (1927.) 
Ann . Trop. Med. & Parasitol ., Vol. XI, p. 92. 


























75 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXV. ARSENIC IN MALIGNANT TERTIAN 

MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 
C. FORSTER COOPER 

AND 

* H. F. CARTER 

From the Liverpool School of Tropical Medicine 

Uhdertaken at the request of the War Office 

(Received for publication 28 March , 1919) 

The observations recorded in this paper were made in order to 
determine the effects of arsenic in malignant tertian malaria. 

All the cases were adult males, infected in either Macedonia or 
West Africa. In every instance a diagnosis of malignant tertian 
malaria was made microscopically, and in all cases trophozoites were 
present in the blood on the day treatment commenced. Blood 
examinations were made daily. 

In the tables and charts : — 

t. = malignant tertian trophozoites, 

c. = malignant tertian gametes (crescents), 

neg. = no parasites found. 

Q.M. = intramuscular injection of quinine bihydrochloride. 

A. = oral administration of Liquor arsenicahs. 

N. = intravenous injection of novarsenobillon. 

Four sets of observations were made as follows : — 

Group A 

A single intravenous injection of novarsenobillon in doses varying 
from gramme 0 45 to gramme 0*9 was given in fourteen cases 



(Nos. 1483 to 1496). The results are recorded in Table I and in 
Charts 1485, 1493, and 1494. 

It will be seen that the treatment had little if any effect on 
trophozoites or on the temperature; in the great majority of cases it 
was found necessary, for clinical reasons, to administer quinine in 
from three to seven days. 


Table I. 

Results of a single intravenous injection of novartenobillon in malignant tertian malaria. 

• W.A. = West Africa. 


Number 

of 

case 

•Place 
of in¬ 
fection 

Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
('in months) 
between 

arrival in 
England 
and 

present 

treatment 

Date 

of 

injection 

Dose 

in 

grammes 

Day of 
injec¬ 
tion 

1st 

day 

after 

2nd 

day 

after 

Paras 

3rd 

day 

after 

tic find 

4th 

day 

after 

ings 

5 th 

day 

after 

6th 

day 

after 

7th 

day 

after 

Sth 

da 1 . 

after 

1483 

W.A. 

. 

1 

0 

13.11.17 

o *45 

t 

t 

f 

tt 




... 


1484 

W.A. 

1 

1 

0 

13.11.17 


t 

c 

c 

t c 

c 

c 

tc 

t ct 


1485 

W.A. 

1 

1 

0 

14.11.17 


t 

t 

t 

t 

t 

t 

t 

t 

t vt 

i486 

W.A. 

1 


0 

14.11.17 


t 

t 

tt 


1 

•• 

... 

... 


1487 

W.A. 

1 


0 

16.11.17 

! >> 

t 

t 

t 

1 

t 

t 

! t ct 

1 


I ... 


1488 


... 


... 

26.11.17 

” 1 

t c 

t c 

1 

c 

t c 

c 

tc 

t 

1 1 c 

t C 

1489 





17.1.18 


t 

t 

neg. 

neg. 

tt 

... 

... 

... 


149° 

W.A. 

1 

1 

0 

26.11.17 

o-6 

t 

t 

tt 

... 

... 


... 

... 

... 

1491 

W.A. 

1 


0 

26.11.17 

51 

t 

t 

t 

neg. 

c 

t 

t 

t 

* t 

1492 

W.A. 

1 

j 

0 

19.II.18 

0-9 

t 

t 

t 

t 

tt 


... 



1493 

W.A. 

1 

1 

0 

19.11.18 


t 

t 

! t 

1 

t 

tt 



... 


1494 

W.A. 

1 

| 

1 

1 

0 

20.11.18 


t 

t 

t 

t 

tt 



... 


1495 

W.A. 

1 | 


0 

23:11.18 

V 

t c 

t 

t 

neg. 

c 

t c 

t 

t c 

t 

1496 

W.A. 

1 i 

1 ! 

0 

1 27.11.18 


t 

neg. 

neg. 

c 

t c 

tt 

... 




t Quinine administered. 































78 


Group B 

Quinine bihydrochloride grains 15 intramuscularly on each of 
two consecutive days only + novarsenobillon gramme o'9 
intravenously on the first day, were given in ten cases (Nos. 1497 to 
1506). The results of this combined treatment are recorded in 
Table II. 

In two of the ten cases treatment was commenced during an 
apyrexial period, in the remaining eight cases the temperature fell 
to normal within two days. Trophozoites disappeared from the 
cutaneous blood within three days. 

Table II. 

Summary of results of administration of quinine bihydrochloride grains 15 intramuscularly on each of two consecutive 
days + novarsenobillon gramme *9 intravenously on the first day, in malignant tertian malaria. 

• S. *■ Salonika. W.A. ** West Africa. 


Number 

of 

case 

•Place 
of in¬ 
fection 

Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 

present 

treatment 

Date 

of 

end of 
treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after first 
injection 

Trophozoites 

disappeared 

from 

cutaneous 
blood 
in — days 
after first 
injection 

Trophozoite 
relapse 
occurred 
in — days 
after first 
injection 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — daw 
alter first 
injection 

H 97 

S. 

26 

1 

1 

9.10.18 

2 

1 

8 

11 

1498 

S. 

39 

3 

3 

14.10.18 

Same day 

1 

*5 


1499 

S. 

27 

2 

1 

14.10.18 

Apyrcxia 

1 

6 

7 

1500 

S. 

25 

2 

1 

22.10.18 

1 

2 

12 

12 

1501 

S. 

28 

2 

1 

24.10.18 

Apyrcxia 

1 

8 

7 

1502 

S. 

18 

2 

1 

26.10.18 

1 

3 

8 

7 

1503 

S. 

2 

2 

2 

28.10.18 

1 

1 

9 

*5 

1504 

W.A. 

1 

1 

0 

2.11.18 

1 

2 ! 

17 

18 

1505 

W.A. 

1 

* 

0 

2.11.1S 

1 

1 

16 

17 

1506 

W.A. 

1 

1 

0 

2.11.18 

1 

1 ! 

*5 

16 


Relafses 

A parasitic relapse occurred in all cases; trophozoites reappeared 
in the blood in six to seventeen days, and febrile relapses occurred in 
seven to eighteen days. 

A comparison of these results with those obtained by the injection 
of quinine bihydrochloride grains 15 intramuscularly on each of two 




79 


consecutive days only, in malignant tertian malaria, recorded in a 
previous paper (1919), shows that the supplementary intravenous 
injection of novarsenobillon had no effect, all the cases relapsing 
within three weeks in both series. 

Group C 

Quinine bihydrochloride grains 15 intramuscularly on each of 
two consecutive days only + Liq. arsenicalis minims 30 by the mouth 
daily (Case 1507). The result of this combined treatment in the 
single case treated is shown in Chart 1507. 

Case 1507 





The temperature fell to normal in two days, and trophozoites 
disappeared from the cutaneous blood in two days after commence¬ 
ment of treatment. Trophozoites reappeared on the fourteenth day 
and a febrile relapse occurred on the fifteenth day. As the condition 
from the fifteenth day onwards was uncontrolled, parasitic febrile 
attacks occurring each day, it was found necessary to abandon the 
treatment on the nineteenth day. From this we conclude that the 
daily administration of Liq. arsenicalis minims 30 had no effect, the 
temporary disappearance of symptoms and parasites being solely 
due to the two intramuscular doses of quinine bihydrochloride. 

Group D 

Liquor arsenicalis minims 30 by the mouth daily for 16 days 
+ Quinine bihydrochloride grains 15 intramuscularly on the first 
and second, eighth and ninth, fifteenth and sixteenth days, were 
given in sixteen cases (Nos. 1508-1523). 









8o 


The result of this combined treatment is recorded in Table III. 
The temperature fell to normal within three days. Trophozoites 
disappeared from the cutaneous blood within three days, except in 
one case (1519) in which they persisted throughout. 

Relapses 

A relapse occurred in every case, trophozoites reappearing in the 
♦ peripheral blood in one to thirty-eight days, average eleven days, 

after cessation of treatment. 


Table III. 

Summary of results of oral administration of Liquor arsenicalis, minims 30 daily for 16 days + quinine bihydrochloride grain# 15 intra¬ 
muscularly on the 1st and 2nd, 8th and 9th, 15th and 16th days, in malignant tertian malaria. 

* S. = Salonika. 


Number 

of 

case 

1 

t 

1 

•Place 
of in¬ 
fection 

[ Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

1 

1 Interval 
(in months) 
between 
arrival in 
England 
and 

present 

treatment 

i 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

1 

1 

Trophozoites 
' disappeared 

1 from 

cutaneous 
blood 

' in — days 
| after 

first dose 

1 

Trophozoite 
relapse 
occurred 
♦ in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Remarks 

1508 


No 

records 


8.1.19 

1 

2 

38 

39 


1509 





2i.1.19 

3 

| 3 

16 


Measles on 18th day. 

1510 





4.2.19 

3 

j 2 

9 


Quinine orally, grs- 4; 1 











10th day. 

1511 





5 . 2 .X 9 

2 

i 2 

8 


Quinine orally, grs. a: 








1 



9th day. 

1512 


1 



6.2.19 

Same day 

1 1 

5 

5 

1 

1513 

1 

i 



11.2.19 

Same day 

i 1 

24 


toi° F. on 12th and 13 


! 






1 



days. Quinine oral 











grs. 45 on 27th day 

15 H 





13.2.19 

Apyrexia 

1 

20 


Quinine orally, grs. 451 











20th day. 

15x5 





25.2.19 

3 

2 

6 

6 

Crescents throughout 








i 



treatment. 

1516 





27.2.19 

1 

1 

5 


Quinine orally, grs. 451 











6th day. 

, 5 I 7 

S. 

6 

4 

0 

27.2.19 

1 

1 

26 

28 


1518 


No 

records 


28.2.19 

1 ! 

1 

5 


Trophozoites on 8 th <UtI 











treatment. Quinn* 











orally, grs. 45 on 











day. 

1519 

s. 

5 

1 

1 

5 - 3-19 

1 

Present 



Quinine orally, grs. y ' 








throughout 



8th day. 

1520 


No 

records 


6.3.19 


1 

9 


Crescents throughout. 







Same day | 




Discharged on 9th Si 

1521 





7 - 3-'9 

Same day 

1 

1 

... 

Quinine orally, grs. 30 i 




l 







7th day. 

1522 





8.3.19 

Apyrexia 

3 

2 

... 

Quioine orally, grs. j- i 

1 










loth day. 

! 5 2 3 

s. 

6 

I 

1 

11.3.19 

2 

3 

10 

... 

Quinine orally, grs. 5: a 

1 










12th day. 





8i 


Table IV. 

Summary of the Treatments. 



Treatment 

Number 
of cases 
observed 

N umber 
of cases 
which 
relapsed 

j Percentage 
j of cases 
which 
| relapsed 

1 

A. 

Novarsenobillon intravenously gm. *45 to gm. ’9 


*4 

100 % 

B. 

Quinine intramuscularly grs. 15 X 2 + Novarseno¬ 
billon intravenously gm. *9 . 

10 

i 

10 

IOO °o 

C. 

Quinine intramuscularly grs. 15 X 2 + Liquor arseni¬ 
calis orally min. 30 daily for 18 days . 

1 

1 


D. 

Quinine intramuscularly grs. 15x6+ Liquor arseni¬ 
calis orally min. 30 daily for 16 days . 

1 

16 

1 

16 

1 '0°% 

Control. Quinine intramuscularly grs. 15 X 2 
(Study XXII) 

29 . ! 

*9 

100% 


CONCLUSIONS* 

1. Novarsenobillon in the doses used is of no value in the 
treatment of malignant tertian malaria. 

2. A combination of arsenic (Novarsenobillon or Liquor 
arsenicalis) with quinine in the doses used is not more effective than 
quinine alone. 


• The results recorded in this paper should be compared with those obtained in simple 
tertian malaria. (Am. Trap. Med. & ParasitoL, Vo!. XII, p. 371.) 




83 


THE INTESTINAL' PROTOZOAL 
INFECTIONS AMONG CONVALESCENT 
DYSENTERICS EXAMINED AT THE 
LIVERPOOL SCHOOL OF TROPICAL 

MEDICINE 

(THIRD REPORT) 

BY 

J. R. MATTHEWS, M.A. 

AND 

A. MALINS SMITH, M.A. 

(Received for publication February, 1919) 

INTRODUCTORY 

The Second Report (1917) on the protozoological examination 
of dysenteric patients received at Liverpool dealt with cases 
admitted to hospital during the period May 1st to December 31st, 
1916. That report contained a full account of the results obtained 
from the examination of 1,713 cases of dysentery and discussed 
various questions connected with the protozoological examination of 
stools. The present report deals with 2,355 cases that have been 
examined for intestinal protozoa during the years 1917 and 1918. 

MATERIAL 

So far as we know, the majority of the cases examined were 
invalided to this country from France. A small number that came 
under our observation had been to the East, but we have no idea 
what proportion of the total number of cases had served on the 
French front only or how many had been in the East before coming 
to France. We are therefore unable to make any comparison of the 
incidence of infection among ‘Eastern 1 and ‘French' cases. We 
believe, however, that the cases here dealt with constitute a fair 
sample of the ‘ convalescent dysenteries * that have returned to this 
country from various fronts during the past two years. 



«4 

ANALYSIS OF RESULTS 

Of the 2,355 cases examined, 1,158, or 49*2 per cent., were 
discovered to be infected with one or more of the intestinal protozoa 
found in man. An analysis of-the 1,158 infected cases is given in 
Table I. The results previously obtained from 1,713 cases (Second 
Report, 1917) are shown for comparison. 


Tabu I. 


Showing incidence of protozoal infections among 2 

,355 convalescent dysenteries 


Number of cases 

Percentage of 

Previous results 

Protozoon 

infected 

total cases 

(Second Report) 




1917 

Entamoeba histolytica . 

! 

306 , 

1 

13-0 

10*9 

E. colt . 

7 C 3 

29-8 

29-4 

E. nana • . 

279 

166 


Giardia intestinalis . 

35 * 

M -9 

18-5 

Cbilomastix mesnili . 

87 

37 


Trichomonas intestinalis . 

i 

11 

'• ! 

„ 


• The percentage of infection with E. nana it based on the last 1,674 cases examined. 
During the early part of the period covered by this report we were not familiar with this proto- 
zoon. 


The results, on the whole, agree fairly closely with those published 
in 1917. The incidence of infection with E. histolytica in the 
present series of cases is somewhat higher than that obtained 
previously, but it will be shown later that the difference can be 
explained. 


DETAILS OF EXAMINATIONS MADE 

In order to appreciate fully the general results shown in Table I, 
it is necessary to know the number of examinations made. This 
matter has so frequently been ignored that we wish to draw attention 
to its importance again. 

The total number of examinations performed was 14,130, which 






«5 


is an average of six per case. This average is obtained from two 
classes of cases, however, and does not give any idea of the actual 
distribution of the total number of examinations. The great 
majority of those cases in which E. histolytica was found received 
more than six examinations each. In fact, 204 E. histolytica cases, 
each examined at least six times, had in all 7,483 examinations, an 
average of 36*7 per case. Apart from these, therefore, there remain 
2,151 cases having 6,647 examinations, which is an average of only 
three per case. The actual distribution is as follows : — 


2,355 cases had at least one examination each. 

i>935 » » two ,, ,, 

1,480 „ „ „ three 

595 *» >> four ,, ,, 

400 ,, ,, ,, five ,, ,, 

3*9 ” ? 5 >> dx ,, ,, 


It is unfortunate that nearly 40 per cent, of our cases should have 
left hospital with only one or two examinations. 

Table II gives the findings of E. histolytica, E. coli and 
G. intestinalis for the number of cases examined at each stage of the 
above analysis. The increase in the number of specific infections 
detected at each succeeding examination is thus indicated. 


Table II. 


I 

Examination 

Number of 
cases 

examined 

E. bist. 
cases 

Per cent. 

1 of total 
j cases 

E. coli i 
cases 

1 

Per cent, 
of total 
cases 

: 

Giardia 

cases 

Per cent, 
of total 
cases 

First 

2,355 

i 8 + 

7-8 

1 

: 366 

* 5-5 

> 211 

1 

9 *o 

Second 

'.935 

244 

IO *4 

551 

23-4 

278 _ 

11*8 

Third 

1,480 

283 

1 ! 2*0 

1 

642 

27*3 

! 309 

* 3 *< 

Fourth 

595 

*95 

12-5 i 

1 

fi 59 

28-0 

! 322 

*37 

Fifth 

1 

400 


127 

666 

28-3 

33° 

14-0 

Sixth 

3 i 9 

30 l 

1 12-8 

67O 

i 28 - 5 

i_ .. 

332 

1 

* 4 * 

Ultimately ... 

i 

306 

! 13*0 

1 

O 

N 

! 

! 29-8 

1 

352 

14-9 




86 


Five infections with E. histolytica detected after the sixth 
examination were found in the following order: two at the seventh, 
one at the eighth, one at the eleventh, and one at the twelfth 
examination.* Thirty-three infections with E. coli not discovered 
until after the sixth examination appeared as follows : — 

Infection detected at the 7th examination in 3 cases. 

,, v 8th ,, 4 ,, 

>> » 9 th * » 2 „ 

„ „ 10th „ 1 case. 

,, nth ,, I ,, 

„ „ 12th „ 2 cases. 

«> >» 1 4 ^ *5 2 ,, 

,, ,, 16 th ,, 2 ,, 

ti it I7th ,y 2 yy 

>» 18th ,, 3 >> 

a 19th ,, 2 ,, 

„ „ 21st „ 1 case, 

n 24th ,, I ,, 

23th yy I yy 

„ ,, 26th „* 2 cases. 

„ „ 28th „ 1 case. 

it 29tll ,, I yy 

a » 45 th „ I 

:t j) 55 1 ^ >> * >> 

Twenty infections with G . intestinalis discovered after the sixth 
examination were found in the following order : — 


Infection detected at the 7th examination in 3 cases. 


it 

»» 

8th 

a 

I case. 

M 


12th 

a 

2 cases. 



15th 

a 

1 case. 

i> 


16th 

a 


it 


17 th 

a 

2 cases. 

it 

>> 

19th 

a 

1 case. 

i) 


22nd 

a 

1 a 

it 

a 

23rd 

a 

* a 

it 


26th 

a 

* it 

H 

» 

27 th 

a 

* a 

it 

?> 

30th 

a 

* a 

it 

?> 

31st 

a 

* a 

ti 

?» 


a 

1 it 

>> 


43 th 

a 

* >> 


>> 

64th 

ty 


# Of course 

very few non* 

‘E. histolytica cases 

were 

examined more than six times. 



87 


INCIDENCE OF INFECTION WITH E. HISTOLYTICA 

It is clear from Table II that if all the cases had received six 
examinations each the percentage of infection would have been much 
higher than that recorded. A full discussion of the real incidence 
of infection with various protozoa was given in our Second Report, 
and we shall refer to the question here very briefly. We gave 
reasons for believing that by a system of six examinations per case 
the incidence of infection with E . histolytica would be about 
18 per cent. At the first examination we had found 6 per cent., 
i.e. the number of positive cases discovered by one examination per 
case was one-third of the number that would have been found had 
each case been examined six times. These calculations have been 
fully confirmed by Mackinnon (1918), who succeeded in examining 
a large series of cases (1,680) six times each. Of this number 
4*3 per cent, were found infected with E. histolytica at the first 
examination. By the sixth examination the percentage had 
increased to 12*4. It will be observed from Table II that 
7*8 per cent, of the present series of cases were found at the first 
examination to be infected with E. histolytica , and we may calculate, 
therefore that about 23 per cent, would have been discovered had 
all the cases been examined six times. The result for the first 
examination is higher than that obtained in our former series, namely 
6 per cent. The difference is due to the inclusion in the present 
series of those infections with E. histolytica whose cysts measure 
less than 10/1 in diameter. These small cysts were not recorded 
before the year 1917, for they were not identified as cysts of the 
pathogenic amoeba. Our early records, therefore, were based on 
the diagnosis of cysts exceeding 10 fi in diameter. Among the 306 
E. histolytica cases included in this report, 110 were infected with 
strains producing cysts under 10 p in diameter, while in 215 
infections cysts larger than 10 /i were found. These figures give 
325 infections in all. In nineteen cases both large and small cysts 
were found. In the 306 E. histolytica cases then, 36 per cent, 
showed an infection with small cysts. This result agrees closely 
with that recorded by Dobell and Jepps (1917), but is considerably 
lower than that reported by Mackinnon (1918). If we had recorded 
the presence of the larger* cysts only the incidence of infection with 



88 


E. histolytica would have been 9'I per cent, instead of 13, and the 
former result would have been similar to that obtained in our 
previous report. 

INCIDENCE OF INFECTION WITH E . COL !, G. INTESTINAL/S 

AND C. MESNE LI 

While we have calculated the probable incidence of infection with 
E. histolytica on a system of six examinations per case, it is possible 
for us to obtain a more accurate idea of the real incidence of 
infection with some of the other intestinal protozoa. For this 
purpose we shall make use of 204 E. histolytica cases that received 
at least six examinations each. Table III shows the number of cases 
infected with E. coli t G. intestinalis and C. mesnili among these 
204 cases. 

Table Ilf. 

Showing number of cases infected '\ith E. coli y G. intestinalis and C. mesnili among 204 
E, histolytica cates examined a large number of times. 


Number of 
cases 

Number of 1 
examina- | 

Entamoeba coli 

Giardia intestinalis 

C. 

mesnili 

examined 

dons per j 
case 

Cases 

Per cent. 

Cases 

Per cent. 

Cases 

1 

Per cent. 

2 °4 . 

. ; 

4 2 

20-6 

*7 

8-3 

7 

3*4 

2 °4 . 

2 1 

62 

30-4 

2 5 

12*2 

10 

4-9 

204. 

! 3 : 

7 * 

38-2 

2 9 

14 ‘ 2 

11 

5*4 

204. 

4 

86 

4 *' 

30 

*47 

12 

5 9 

204. 

5 : 

9 * 

i 446 

I 

35 

« 7 -« 

13 

64 

204 . 

i 

i 6 

93 

1 45-5 

3 6 

176 

! 4 

6’8 

204. 

1 

Average of 

367 

122 

1 59-8 

! 54 

26-4 

29 

! 4 * 2 


Table III shows clearly that the increase in the number of 
infected cases foupd is the direct result of the increasing number of 
examinations made upon each case. The number of G. intestinalis 
cases detected at the first examination was 17, or 8 3 per cent., which 
agrees closely with the result for the whole series of cases (see 
Table II). Ultimately, however, 264 per cent, of the 204 





8 9 


% 


E. histolytica cases were discovered to have Giardia, a result which 
agrees very closely with an observation made in our Second Report 
(see Table V of that report), where, among no £. histolytica cases 
26*3 per cent, were ultimately found to be infected with Giardia. It 
is probable, therefore, that if the whole series of 2,355 cases had been 
examined many times each the incidence of infection with Giardia 
would have been found to approximate to 30 per cent. 

Similar observations may be made regarding the incidence of 
infection with E. coli . Table III shows that the percentage increases 
from 20*6 at the first examination to 59*8 per cent, ultimately. 
There is reason to believe, however, that E . coli occurs more 
commonly among E. histolytica cases than in a random selection of 
cases. At the third examination of the 204 cases considered in 
Table III it will be seen that 38*2 per cent, were infected with 
E. coli, while in the total series of 2,355 cases with an average of 
three examinations per case only 29*8 per cent, were found infected. 
If we suppose that the percentage figures for E. coli given in 
Table III are approximately 10 per cent, higher than they would be 
for an unselected series of cases, we obtain 50 per cent, as the 
probable real incidence of infection with this amoeba. This agrees 
with the conclusion reached in our former report. 

Particulars have also been given in Table III for C . tnesnili. 
Among the 204 selected cases, 14*2 per cent, were ultimately found 
to be infected with this flagellate. In our former report we 
concluded that ‘ it was not improbable that the real incidence of 
infection was 12 to 15 per cent./ and the present data give further 
evidence in support of that conclusion. 


ORGANISMS OTHER THAN PROTOZOA 

The commonest helminthic infection was Trichuris trichiura, the 
eggs of this worm being found in 110 cases (4*7 per cent.) Ascaris 
lumbrtcotdes was recorded in ten cases. Eggs of Taenia sp. were 
observed in three cases, Oxyuris vermicularis in two cases and 
Strongyloides sp. on one occasion. Iodine cysts (Wenyon) were 
found in forty-one cases (1*7 per cent.). 



9 ° 


4 


r£sum 6 of results for all the dysentery cases 

EXAMINED AT LIVERPOOL SINCE MAY, 1916 

It may be useful here to bring together the results of all the work 
done on the examination of dysenteric patients at the Liverpool 
School of Tropical Medicine since May, 1916. Altogether 4,068 
cases have been dealt with, and 23,024 microscopic examinations of 
the stools performed. Protozoal infections were discovered in 1,976 
cases (48*5 per cent.), an analysis of the infected cases being shown 
in Table IV. 

Table IV. 


Number of cases Percentage of total 
infected 1 cases examined 

K. histolytica . ..J 404 

E. coli .■ l,2oX 

i 

E. nana • ... ... ... ... ... ...i 279 

C. intestinal is ... ... ... ... ... ... 669 

C. mesni/i ... ... ... ... ... ... 148 

T. inteststtalis . .. 29 

• Based on last 1,674 cases examined. 


REFERENCES 

Carter, Mackinnon, Matthews, and Smith (1917). Protozoological Investigation of cases of 
dysentery conducted at the Liverpool School of Tropical Medicine (Second Report). 
Annals. Trop. Med. Parasitol ., Vol. XI. pp. 27-68. 

Dobell and Jepps (1917). On the Three Common Intestinal Entamoebae of Man. Frit. Med. 
Journ ., May 12, pp. 607-612. 

Mac kin non (1918). Notes on the Intestinal Protozoal Infections of 1,683 mcn examined at 
the University War Hospital, Southampton. Lancet , September 21, pp. 386-389. 


12*1 

297 

166 

164 

3-6 

07 


Protoroon 













9 ' 


THE SPREAD AND INCIDENCE OF IN¬ 
TESTINAL PROTOZOAL INFECTIONS IN 
THE POPULATION OF GREAT BRITAIN 

IV. ASYLUM PATIENTS 
V. UNIVERSITY AND SCHOOL CADETS 

BY 

J. R. MATTHEWS, M.A. 

AND 

A. MALINS SMITH, M.A. 

(Received for publication 18 March , 1919) 


IV. ASYLUM PATIENTS 


In continuation of our general investigation of the intestinal 
protozoa of the population of this country (Matthews and Smith, 
1919) we examined (June, 1918 to February, 1919) specimens of 
faeces from two hundred and seven patients of the Lancashire 
County Asylum, Whittingham. This we were enabled to do by the 
kind permission of Dr. J. F. Gemmel, who also helped us by 
furnishing information as to the age, length of residence, and state 
of health of the patients whose stools were examined. The patients 
were all males, and ranged in age from 17 to 87, the average age 
being 48. The protozoal infections found among these patients, as 
the result of a single examination, were as follows: — 


Entamoeba histolytica 

E . coli . 

E . nana . 

Giardia intestinalis 
Chilomastix mesnili 


97 per cent. 

45*9 

121 

3*4 

23 ‘ 2 


Besides these protozoal infections we found the eggs of Trichuris 
trichiura in 3*4 per cent, of the cases, and those of Oxyuris 
vermicularis in 1*9 per cent. 


4 




9 2 


In Table I the above figures are given together with the results 
for other sections of the population examined by us and also for 
dysenteric and non-dysenteric returned soldiers. All the figures, 
with the exceptions stated, are the results of a single examination. 



Never out of Britain. 

1 

Returned Soldiers 


Asylum 

Patients 

Adult 

civilians 

Armv j 
Recruits | 

1 

1 

1 

Cliildren 

! 

Dysenteric 

Convalescents 

1 

Non- 

dysenteric 

Convalescents 

No. examined 

207 


1098 

548 I 

40^8 

450 

E. histolytica 

! 

97 

p 5 

5-6 

1 

i-8 

7 *° 

6-4 

F. coli . 

45*9 

67 

18-2 

1 

1 pi 

15*2 

142 

F. natia . ... 

12 1 

2*4 

5‘5 

27 

— 

— 

G. intestinalis 

3*4 

6 -o 

7 *o 

mi 

9‘9 

6-o* 

C. mcsv.ili ... 

1 2 yz 

1 

i*5 

2 cases 

1 

; p8 

3-6 

(3 exams, per 
case) 

2*0 

(2 exams, per 
case) 


We have stated in previous papers that at least five hundred cases 
should be examined before reliable figures can be obtained as to the 
true incidence of the various protozoal infections in any population. 
Unfortunately in the case* of Asylum patients we have not been able 
to examine more than two hundred and seven, a number too small 
to give reliable comparative figures. We can therefore only indicate 
one or two outstanding features which we think are significant, 
without considering any of the results as definitely established. 

The first striking feature is the high percentage of almost all the 
infections among the Asylum patients. Infections with all the 
protozoa but Giardia intestinalis are more numerous than in any 
other group we have examined, not excepting those who have been 
to tropical and sub-tropical countries. In spite of the small numbers 
examined this seems to represent some real difference in the present 
group, and it may perhaps be suggested that the infections spread 
more rapidly and more widely within this isolated population 
because of the well-known lack of cleanly and careful personal habits 
among the insane. 

* In Matthews and Smith (1919) Army Recruits, Tabic V, this value is given in mistake 
as io*8. The present figure is correct. 






93 


A further noteworthy result is shown in the figures for Giardia 
intestinalis infections. These are a complete exception to all the 
remaining results, for not only are they not higher than in the other 
population groups, but they are considerably lower than in any 
other group. This fact seems to us to fit in with the suggestion we 
put forward in our paper on the infection of children, namely that 
G. intestinalis is mainly a parasite of children and becomes rarer in 
older people. The present group of Asylum patients with average 
age of 48 years is the oldest section of the population which we have 
examined, and it appears significant that, while all other infections 
are so common, this one should be rare. It is particularly striking 
to find G. intestinalis comparatively uncommon, since infections with 
C. mesnili , the other common flagellate of the human intestine, are 
very numerous, being many times more common than in any other 
group we have examined. Both flagellates were in every case found 
in the encysted state in the faeces, and we cannot suggest any cause 
of the remarkably different incidence of the two, other than that 
G . intestinalis tends to disappear in older people, while C. mesnili 
does not. 

We hoped at the outset of the investigation to obtain some 
evidence as to the effect of continued residence in the institution 
upon the incidence of infection and also to find out whether those 
numerous patients, who suffer from colitis, were more or less heavily 
infected than the remainder, but the number of the patients examined 
has not been sufficient to give us any evidence upon these points. 

We tender our hearty thanks to Dr. J. F. Gemmel, the Medical 
Superintendent of the Asylum, and also to Sergt. Fann, R.A.M.C., 
for his efficient help in the collection of the specimens. 


SUMMARY 


Two hundred and seven Asylum patients have been examined for 
intestinal protozoa. All the protozoa, but G. intestinalis , were 
found more commonly than in any other population group we have 
examined. G. intestinalis , on the other hand, was less common than 
in any other group. 



94 

V. UNIVERSITY AND SCHOOL CADETS 

We began in April, 1918, an examination of the stools of youths 
in the Liverpool University Platoon of the Manchester University 
Officers* Training Corps, and also of the older boys.in a high-class 
secondary school near Liverpool. 

We examined in all only forty-one such persons, all males and 
ranging in age from 15 to 19 years. As it seems now very 
improbable that the investigation will be continued, we give without 
comment the results so far obtained. None of those reported on had 
ever been out of this country. 

No. examined 

E. histolytica 
E. coli 
E. nana 
G. intestinalis 
C.mesnili 

The results are from a single examination. Our object was to 
examine a number of youths from a higher social class and from 
better homes than the Army recruits whom we had previously 
reported on. We can only say that infections of the various protozoa 
exist among such persons. 


41 

1 case 
II cases 
1 case 

. 1 „ 


REFERENCE 


Matthews, J. R., and Smith, A. Malins (1919). The Spread and Incidence of Intestinal 
Protozoal Infections in the Population of Great Britain. I. Civilians in Liverpool 
Royal Infirmary. II. Army Recruits. III. Children. Anti. Trop. Med. & Parantoi. 
Vol. XII, p. 349 - 





95 


AWARD OF THE MARY KINGSLEY 

MEDAL 


At a meeting of the Committee of the School, held on the 
ioth March, 1919, it was resolved to award this medal to 
J. W. S. Macfie, M.A., M.B., D.Sc., in recognition of his 
distinguished scientific work in Tropical Medicine. 

Dr. Macfie was appointed to the West African Medical Staff, 
Northern Nigeria, in June, 1910. In 1913 he was seconded to the 
Medical Research Institute, Lagos, as bacteriologist, and during that 
year took part in the Yellow Fever investigation in West Africa. 
In 1914 he became pathologist at Accra, and in March, 1917, was 
seconded to the Liverpool School of Tropical Medicine to undertake 
research on the treatment of malaria. 

During the last nine years Dr. Macfie has published many 
valuable contributions to science, notably on trypanosomiasis in man 
and domestic stock. 

The medal was presented to him by the Chairman of the School 
at a luncheon given on the 12th April, 1919. 

At the commencement of this number will be found a list of the 
medalists. 







97 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXVI. THE ACTION OF ARSENIC AND OF 
QUININE ON QUARTAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 

AND 

J. W. S. MACFIE 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
(Received for publication 1 May, 1919) 

Two intravenous injections of novarsenobillon 0*9 gramme were 
given at an interval of approximately a week in two cases (Nos. 1524 
and 1525). The results, which are given in the temperature charts, 
show that the treatment had no appreciable effect either on the 
temperature or on the parasites. After it had been ascertained that 
novarsenobillon was ineffective, an intramuscular injection of quinine 
bihydrochloride grains 15 on each of two consecutive days was given 
in each of the cases. From the charts it will be seen that in neither 
case did the injections cause the disappearance of the parasites, 
trophozoites and gametes persisting. In one case (1524) the 
temperature was controlled, in the other (1525) it was not. 

T. = quartan trophozoites or schizonts. 

G. = quartan gametes. 

Neg. = no parasites found. 

N. = intravenous injection of novarsenobillon. 

Q.M. = intramuscular injection of quinine bihydrochloride. 

Q.O. = oral administration of quinine sulphate. 

A comparison with the results given in Studies XVI, XXV, and 
XXVII shows a remarkable difference in the action of novarseno¬ 
billon and quinine on the three species of malaria parasites. 



98 


Case 1524 


Will 

■nil 

mBBiBiirimi 


Mill 

■HI 


111 — 


P^IJ^DBBBBHHM 


IBM 


ijSIIKfii 


iiaiaBB BB BriyiaayiyiQaMa 

■MMMria^MMHBMnnBH 


iiOiOUK 
-1311- 


|CIESIllQ]£SE!3ESE!SLiJliJE!IEQC!inEIEIC9anCIDiEIDOir9IIllflCflDl 

b^bbbbbbbbbbbbb^bbbbb^^bbbbI 

BBBBBBBBBBBBBBBBBBBBBBBBBIBBBBI 
BBBBBBBBBBBBBBBBIBBBBBBBBBBBBBI 


lBIB t lBBB BiBBHBB»BBnH BBHHBHHBHBI 

K BBiM gjBlHgPtaiBlgBB——i 


I333E1 




















99 

NOVARSENOBILLON 


The action of novarsenobillon on Plasmodium vivax is marked. 
In this infection its action is even more rapid and efficient than that 
of quinine, a single intravenous injection of 0 9 gramme causing the 
disappearance from the cutaneous blood of all stages of the parasites 
within twenty-four hours. In the case of P. falciparum and 
P. malariae novarsenobillon in the same dosage has no appreciable 
effect on the temperature or on the parasites. 


QUININE 

An intramuscular injection of the bihydrochloride of quinine 
grains 15 on each of two consecutive days only, exerts in the case 
of P . vivax a constant and rapid effect both on the temperature and 
the parasites; in the case of P. falciparum the action on the tempera¬ 
ture and trophozoites is also well defined, though relapses occur more 
quickly than in the case of P. vivax , whilst in the two cases of 
P . malariae treated in the same way there is little if any effect on the 
parasites, but in one of the two cases the temperature was controlled. 


REFERENCES 


Stephens, J. W. W., Yorke, W., Blacklock, B., Macfie, J. W. S., Cooper, C. F., and 
Carter, H. F. (1918 and 1919). Ann. Trop . Med. & Parasitol ., Vol. XII, p. 21 1 , 
and Vol. XIII, pp. 75 and 101. 




IOI 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXVII. INTRAVENOUS INJECTIONS OF 
NOVARSENOBILLON AND INTRAMUSCULAR 
INJECTIONS OF QUININE BIHYDRO¬ 
CHLORIDE IN SIMPLE TERTIAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 

AND 

J. W. S. MACFIE 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Oifice 
{Received for publication 5 May, 1919) 

The observations made by us can be divided into three groups. 

GROUP I. A single intravenous injection of novarsenobillon 
0 9 gramme on the first day with an intramuscular injection of 
quinine bihydrochloride grains 15 on the first and second days of 
treatment. 

Group II. An intravenous injection of novarsenobillon 
o - 9 gramme on the first, eighth, and fifteenth days, with intra¬ 
muscular injections of quinine bihydrochloride grains 15 on the 
first and second, eighth and ninth, and fifteenth and sixteenth days 
of treatment. 

GROUP III (control series). An injection of quinine bihydro¬ 
chloride grains 15 on each of two consecutive days only. 



102 


GROUP I (Cates 1526 to 1566) 

In nine of the forty-one cases (Table I) treatment was commenced 
during an apyrexial period. In thirty cases the temperature fell to 
normal within one day, in one case (1533) in two days. In one case 
pneumonia developed on the second day of treatment. 

In all the cases parasites disappeared from the blood in one day. 

Relapses 

Thirteen of the forty-one cases (32 per cent.) relapsed within the 
sixty-days* observation period. Parasites reappeared in thirteen to 
forty-eight days (average twenty-nine days), and febrile relapses 
occurred in fourteen to forty-eight days. 

Two of the cases (1543 and 1551) which did not relapse within 
the sixty-day observation period did so on the sixty-fourth and 
sixty-fifth days respectively. 


GROUP II (Cases 1567 to 1578) 

In five of the twelve cases (Table II) treatment was commenced 
during an apyrexial period; in the remaining seven the temperature 
fell to normal within two days. 

In all cases parasites disappeared from the cutaneous blood in 
one day. 

Relapses 

One case relapsed on the last, i.e. the sixtieth day of the post- 
treatment observation period. 

In one case (1578) the observation period was less than sixty 
days, viz., forty-five days; consequently the minimum number of 
relapses, those actually observed, is 8 per cent, and the possible 
maximum number 17 per cent. 

One case (1574) which did not relapse within the sixty-day 
observation period did so on the sixty-eighth day. 



io3 


Table I. 

mary of results of a single intravenous injection of novarsenobillon gramme 0*9 on the 1st day, with an intramuscular injection of quinine 
bihydrochloride grains 15 on the 1st and 2nd days of treatment, in simple tertian malaria. 

• E.A. = East Africa. F. = France. I. = India. M. = Mesopotamia. S. = Salonika. 


•Place 
of in¬ 
fection 

Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 
not 
relapse 

Remarks. 

S. 

28 

10 

9 

3.10.18 

Same day 

1 



*27 


S. 

>3 

9 

9 

3.10.18 

1 

1 


... 

62 


S. 

*5 


... 

3.10.18 

1 

1 



>24 


S. 

21 

3 

2 

8.10.18 

1 

1 



106 


S. 

M 

4 

2 

9.10.18 

Same day 

1 


... 

72 


I. 

16 

7 

3 

9.10.18 

Same day 

1 


... 

r 


S. 

28 

5 

5 

10.10.18 

Apyrexia 




6< 


E.A. 

IS 

9 

7 

10.10.18 

2 

1 

35 

... 

•• 

Quinine orally, grs. 45 
on 44th day. 

E.A. 

8 

8 

7 

11.10.18 

Apyrexia 

1 

27 

29 

•• 


S. 

16 

5 

4 

11.10.18 

1 

1 

... 

... 

103 


E.A. 

1 

>5 

9 

7 

11.10.18 

Same day 

1 

39 

39 

... 


F. 


13 

*3 

11.10.18 

Same day 

1 

... 

... 

113 


... 

... 



11.10.18 

Same day 

1 



7 ° 


E.A. 

10 

9 

7 

12.10.18 

Apyrexia 

1 

... 


80 


S. 

28 

2 

2 

13.10.18 

Same day 

1 


... 

68 


E.A. 

26 

7 

1 

13.10.18 

Apyrexia 

1 

... 

... 

69 


S. 

42 


1 

15.10.18 

Apyrexia 

1 

... 

... 

118 


S. 

H 

2 

2 

16.10.18 

1 

1 

1 

I 

... 

63 

Relapsed on 64th day. 

S. 

14 

10 

4 

17.10.18 

1 

1 

28 

... 

... 

Quinine orally, grs. 45 
on 37th day. 

M. 

28 

27 

4 

20.10.18 

1 

1 

1 

... 

116 


S. 

26 

7 

7 

20.10.18 

Apyrexia 

1 

... 

... 

100 






Table I —Continued 


Number 

of 

case. 

•Place 
of in¬ 
fection 

Interval 
(m months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months] 
between 
arrival in 
England 
and 

present 

treatment 

i 

! Date 
of end 

1 of 

treatment 

Tempera- 
| ture fell 
. to normal 
i in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 

I not 

relapse 

Re marts. 

*547 

S. 

7 

5 

4 

22.10.18 

1 

1 

... 


IOI 


1548 

S. 

28 

2 

2 

23.10.18 

Same day 1 



in • 


•549 

S. 

11 

4 

3 

24.10.18 

Same day 

1 



97 


! 55 ° 

S. 

12 

t 

0 

25.10.18 

Same day 




97 


1 55 1 

S. 

27 

7 

6 

25.10.18 

1 

1 



64 

Relapsed on 65th is 

*552 

... 

... 

... 

... 

25.10.18 

Apyrexia 

1 

*3 

14 

... 


*553 

S. 

28 

4 

3 

25.10.18 

Apyrexia 


20 

21 

... 


>554 

s. 

7 

4 

2 

27.10.18 

X 


... 


96 


>555 

s. 

>3 

5 

4 

27.10.18 

Same day 

1 

... 


63 


1556 

! E.A. 

! 

10 

7 

6 

27.10.18 

Apyrexia 

1 

43 


... 

Quinine orally, gn. ] 
on 44th day. 

>557 

E.A. 

9 

7 

5 

27.10.18 

1 

1 

18 

18 

... 


1558 

s. 

1 

23 

2 

2 

27.10.18 

Same da y 



i 

82 


>559 

S. 

27 

2 

2 

27.10.18 



48 

48 j 

... 


1560 

s. 

... 

2 

1 

29.10.18 

I 


... 


9 * 


I56I 

s. 

29 

8 

8 

i 

31.10.18 

1 I 

1 

3 i 


... 

Quinine orally, gn.; 
on 32nd day. 

1562 

s. 

24 

6 

5 1 

1.11.18 

1 ! 

1 

3 ° 

3 o 

... 


1563 

I. 

0 

3 

z 1 

10.11.18 

* 

> i 

... 

... 

69 


>564 

E.A. 

*4 

8 

7 

12.11.18 

Same day | 

1 

1 

*5 

... 

... 

Quinine orally, grs 4 
on 27th day. 

>565 ! 

S. 

8 

8 

2 ; 

1 

I 

26.11.18 

1 

1 

*7 



Quinine orally, gn. 3 
on 18th day. 

I566 

S. 

1 

2 

2 

29.11.18 

1 

! 

1 

1 

1 



64 

Pneumonia on 3 i 
day of treat 

-4 







ic >5 


Table II. 


Summary of results of an intravenous injection of novarsenobillon gramme 0*9 on the ist, 8th and 15th days, with intramuscular injections 
of quinine bihydrochloride grains 15 on the ist and 2nd, 8th and 9th, 15th and 16th days of treatment, in simple tertian malaria. 

# E.A. = East Africa. I. = India. S. = Salonika. 


imber 

of 

ase. 

•Place 
of in¬ 
fection 

Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infectecf 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 
not 
relapse 

Remarks. 

5*7 

E.A. 

*4 


9 

1.12.18 

Apyrexia 

1 

60 



Quinine orally, grs. 30 
on 61st day. 

568 

E.A. 

12 


9 

1.12.18 

Apyrexia 




65 


569 

S. 

25 


2 

1.12.18 

2 

1 



7 i * 


570 

j s. 

2 


4 

4.12.18 

2 

1 



65 


57 i 

s. 

13 


4 

4.12.18 

1 

1 

... • 


65 


572 

s. 

28 

1 

2 

6.12.18 

Apyrexia 

1 



66 


573 

s. 

18 


7 

8.12.18 

Apyrexia 

1 



82 

100*2° F. on last day 
of treatment 

574 

I. 

12 


5 

11.12.18 

1 

1 



67 

Relapsed on 68th day. 

575 

s. 

6 

3 

2 

11.12.18 1 

1 2 

1 



76 


576 

s. 

31 

3 

3 

12.12.18 

« 

1 



60 


577 

s. 

24 

*5 

6 

14.12.18 

Same day 

1 



76 


578 

s. 

l 7 

8 

7 

15.12.18 

Apyrexia 

1 



45 














io6 


Tabii III. 


Summary of results of an intramuscular injection of quinine bihydrochloride grains 15, on each of two days only, in simple tertian malaria. 
# E.A.= East Africa. I.= India. M. =3 Mesopotamia. S.= Salonika. 


Number 

of 

case. 

•Place 
of in¬ 
fection 

Interval 
(in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

1 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 
not 
relapse 

Remarks. 

1579 

S. 

29 

3 

3 

1.11.18 

Same day 


29 

... 

... 

Quinine on 32nd dji 

00 

0 

S. 

<4 

6 

5 

2.11.18 

Apyrexia 

2 

20 



Quinine orally, gn . 51 
on 22nd day. 

1581 

S. 

3 ° 

2 

2 

5.11.18 

2 

2 

20 

21 

... 


1582 

M. 

29 

29 

5 

5.11.18 

1 

3 

20 

23 

... 


1583 

S. 

22 

1 

6 

5 

6.11.18 

« 

. 



77 


00 

w-i 

S. 


6 

5 

6.11.18 

Apyrexia 

. ! 

1 

22 

24 



» 5*5 

S. 

<5 

4 

3 

9.11.18 

Same day 

1 

22 

20 

... 


I586 

S. 

29 

3 

2 

9.11.18 

Same day 


... 

... 

80 


* 5*7 

S. 

8 

8 

2 

9.11.18 

Apyrexia 

1 ! 

<3 

<4 



1588 

E.A. 

10 

9 

3 

10.11.18 

1 

« ; 

18 

18 



■ 5 8 9 

- 

! 

... 

... 

26.11.18 

Same day 

1 1 



37 


1590 

E.A. 

37 

... 

5 

1.12.18 

Same day 

1 1 

18 

18 

... 


1 59 1 

S. 

16 

4 

3 

11.12.18 

Same day 


10 


... 

Quinine orally, gn. ? 
on nth diy. 

* 59 * 

S. 

27 

2 

1 

14.12.18 

Same day 

1 1 

... 

! 

60 


1593 

E.A. 

10 

6 

0 

15.12.18 

2 i 

1 

1 

30 

sffil 

... 

Quinine orally, gn 52 
on 32nd day. 

<594 

I. 1 

34 

7 

2 

21.12.18 

* 

2 

5 < 

jSH 

... 


<595 

S. 

4 

1 

1 

9<-<9 

Same day 

1 

21 


... 

Discharged on 25 i 
day. 

l ^ 
* 


... 

... 


9 - 3 * <9 

Same day 

1 

1 

<3 

<5 

... 








Table IV. 


Comparison of curative results obtained from the different treatments. 


Treatment 

Number 
of cases 
observed 
after 

treatment 

1 

. Number 
Number of cases 
of cases not 

which relapsing 

relapsed but 

Percentage of cases 
which relapsed 

within 

60 days t 

observed 
for less than 
60 days 

Minimum 

Maximum 

Group I. Novarsenobillon gm. 

o*9 intravenously on 
the ist day, with 
quinine bihydrochl., 
grs. 15 intramuscu¬ 
larly on the i st and 
2nd days . 

4 i 

1 

I 

*3 

0 

3 *% 

j*% 

Group II. Novarsenobillon gm. 

0*9, intravenously on 
the ist, 8th and i$th 
days, with quinine 
bihydrochl. grs. 15 
intramuscularly on 
the 1st and 2nd, 8th 
and 9th, 15th and 
16th days . 

12 

1 

X 

1 

8% 

■ 7 % 

Group III. Quinine bihydrochl. 

grs. 15 on each of 
two days only (Con¬ 
trol Series). 

18 

*4 

i 

1 

78% 

OO 

O'' 

•Study XVI. Novarsenobillon gm. 
(Table III.) 0*9 intravenously 

(single injection) 

21 

*9 

1 

9 °% 

95 % 


Vide Reference. 





io8 

GROUP III (Cues 1579 to 1596) 

In three of the eighteen cases (Table III) treatment was 
commenced during an apyrexial period. In the remaining fifteen, 
the temperature fell to normal within two days. 

In seventeen cases parasites disappeared from the cutaneous 
blood within two days, in one case within three days. 

Relapses 

Fourteen of the eighteen cases relapsed within the sixty-days’ 
observation period. Parasites reappeared in ten to fifty-one days 
(average twenty-two days) and febrile relapses occurred in fourteen 
to fifty-two days. 

In one case (1589) the observation period was less than sixty 
days, viz., thirty-seven days; consequently the minimum number of 
relapses is fourteen (78 per cent.) and the possible maximum fifteen 
(83 per cent.). 


CONCLUSION 

A combination of novarsenobillon and quinine is more effective 
than either novarsenobillon or quinine alone. 


REFERENCE 

Stephens, J. W. W., Yorke, W., Blacklock, B., Macne, J. W. S., Cooper, C F., and 
Carter, H. F. Ann , Trop, Med, & Parasitol VoL XII, p. 215. 



FILAR1A PERTENUE, n.sp,, 
PROVOQUANT UNE ETERMO- 
FILARIOSE CHELOIDIFORME CHEZ 
CEPHALOPHUS STLFICULTOR 


PAR 

J. RODHAIN 
(Plates II and III) 

(Received for publication 8 April, 1919) 

Le nematode dont nous donnons ici une description, a ete 
recueilli chez la grande Cephalope: Cephalophus sylvicultor a 
Bambili sur l’Uele au Congo Beige. 

L’antilope, qu’on nous apporta tuee, portait au haut des cuisses 
deux taches glabres au niveau desquelles la peau noire, legerement 
saillante et luisante, avait pris l’aspect qu’ont les grosses cheloi'des 
des tatouages negres. Ces plaques cutanees, allongees, ayant 
1 et l centimetre de large sur plus de 5 cen. de long, occupaient 
une position sensiblement symetrique. Toutes deux legerement 
incurvees se rapprochaient en haut et en dedans de la base 
d’insertion de la queue, et s’en ecartaient obliquement en bas et en 
dehors. 

Un examen microscopique rapide des raclures prelevees au scalpel 
dans le tissu indure, montra a cote de debris d’helminthes des 
embryons filariens. Des fragments de la lesion vermineuse furent 
fixes <t l’alcool et au formol pour etude ulterieure. Celle-ci 
commencee k Leopoldville au d6but de 1915 a pu etre complete 
r^cemment au laboratoire de M. E. Roubaud k l’Institut Pasteur a 
Paris. 

DESCRIPTION DE LA L&SION ANATOMIQUE 

Macroscopiquement , la pseudoch 61 oide presente a la section 
l’aspect harolac6 du tissu fibreux, mais parseme de nombreux points 
et trainees jaunatres. Ces dernieres correspondent aux trajets 
tortueux des vers. Elle s’etend en profondeur, de l’^piderme 




IOI 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXVII. INTRAVENOUS INJECTIONS OF 
NOVARSENOBILLON AND INTRAMUSCULAR 
INJECTIONS OF QUININE BIHYDRO¬ 
CHLORIDE IN SIMPLE TERTIAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 

AND 

J. W. S. MACFIE 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
(Received for publication 5 May, 1919) 

The observations made by us can be divided into three groups. 

GROUP I. A single intravenous injection of novarsenobillon 
o'9 gramme on the first day with an intramuscular injection of 
quinine bihydrochloride grains 15 on the first and second days of 
treatment. 

GROUP II. An intravenous injection of novarsenobillon 
0 9 gramme on the first, eighth, and fifteenth days, with intra¬ 
muscular injections of quinine bihydrochloride grains 15 on the 
first and second, eighth and ninth, and fifteenth and sixteenth days 
of treatment. 

GROUP III (control series). An injection of quinine bihydro¬ 
chloride grains 15 on each of two consecutive days only. 



IIO 


jusqu’au contact des muscles fessiers, atteignant i centimetre de plus 
grande £paisseur. 

Les coupes microscopiques montrent que l’epaisissement et 
l'induration de la peau sont dus a PJiypertrophie sclereuse du derme 
provoqu6e par Tirritation chronique causee par les nematodes qui 
habitent le chorion. Les helminthes, plies et replies parfois coudes 
sur eux-memes dans le tissu conjonctif fibreux, y determinent une 
reaction inflammatoire du type essentiellement chronique. Celle-ci 
se reconnait au voisinage immediat des parasites. Ils sont entoures 
d'une veritable gangue de pus epais constitue par les Elements 
conjonctifs dissoci^s et necroses entre lesquels se sont infiltres des 
phagocytes mononucleaires, petits et grands lymphocytes entremeles 
de macrophages et plasmocytes. La ou les fragments de nematodes 
se voisinent, les gangues de pus confluent, il se forme dans le tractus 
fibreux des noyaux de tissu infiltre, friable, veritable debut de petites 
cavites (Planche II, figs, i et 2). 

En dehors des parasites, la trame conjonctivale du chorion 
r^agit par la formation de tissu fibreux dense dont on peut observer 
les divers stades de differenciation. L’hypertrophie sclereuse qui 
en resulte, entraine secondairement l’an&nie de la couche papillaire 
du derme, Tamincissement de Tepiderme, Tatrophie et la disparition 
des follicules pileux et des glandes sebacees, la reduction des 
glandes sudoripares. L'epiderme au niveau de la lesion filarienne 
prend Faspect luisant cicatriciel. 

Les parasites siegent dans les couches profondes du derme et 
ne p^netrent que rarement dans les couches superieures. Dans 
celles-ci se produit une stase lymphatique que traduit la dilatation 
du r^seau sous-papillaire. Au niveau de la cheloide, le tissu 
cellulaire souscutan^ a disparu et le derme hypertrophie, s’etend 
jusqu’a la couche musculaire des fessiers auxquels la tumeur est 
adherente. Les filaires, du moins leurs extremites, se deplacent 
dans les tissus qu’ils irritent. Leurs anciens trajets se reconnaissent 
sur les coupes, aux trainees de pus caracteristiques, dont les 
canalicules vides de parasites sont bientot envahis par les 416 ments 
migrateurs de leurs parois meme. Ult^rieurement ces trajets sont 
combles par du tissu de neoformation derive des fibroblastes suivant 
un veritable processus cicatriciel. 

Ces 16 sions sont en plus d’un point comparables aux nodules 



Ill 


fibromateux produits par VOnchocerca volvulus chez Thomme. 
Mais ici les parasites siegent dans les canaux ou les ganglions 
lymphatiques. Ils sont moins nombreux et plus immobiles dans 
leurs tumeurs. Leur localisation fibreuse est plus stricte. Plus 
volumineux, ils cr6ent dans leurs fibromes de v£ritables cavit6s dans 
lesquelles les extremites de males et de femelles voisinent. Dans la 
cheloide filarienne de Cephalophus sylvicultor , ces cavit6s ont leur 
homologue dans les noyaux de pus ou les fragments de vers sont 
rapproch^s et dehors desquelles nous avons pu aussi isoler des 
extremites d’helminthes. 

DESCRIPTION DES FI LAI RES 

• 

En dissociant sous le binoculaire des coupes epaisses de la 
cheloide vermineuse, nous en avons extrait des fragments de vers, 
parmi lesquels les extremites anterieures et post6rieures de male et 
de femelle dont Tetude a permis la classification zoologique du 
parasite. II s’agit d’une des Filariinae que nous rattachons au genre 
Filaria tel que celui-ci persiste actuellement. 

Le corps est filiforme, attenue k ses deux extremites dont la 
posterieure est plus amincieque Tanterieure. La cuticule est lisse,' 
presentant au niveau des courbures ou des coudures du ver de 
nombreux plissements et montrant aux forts grossissements sous un 
eclairage favorable une tres fine striation transversale. La tete est 
plus ou moins reguli£rement arrondie, parfois tres legfctement 
aciculee vers son sommet, au niveau de la bouche. Celle-ci chez 
certains exemplaires apparait proeminente mais est toujours sans 
levres distinctes ni papilles visibles.* L’oesophage est droit sans 
dilatations. L’intestin des son origine est plus large que la partie 
terminale de l'oesophage. 

Mdle . Longueur inconnue.t Plus etroit que la femelle, il ne 
mesure que 72 fi a 78^ de largeur en son plein d6veloppement. Son 
extremite ant6rieure regulierement retrecie n’atteint plus que 61 a* 
de large immediatement en arriere de la bouche. L'extremite 

• Lorsqu’ au cours des manipulations de la preparation des parasites, Textremite 
cephalique est comprimee, la partie buccale devient nettement saillante, et il apparait une 
indication de demarcation entre la tete et le corps. Chez les individus non deform^s cette 
delimitation n’est pas visible. 

t Le plus long fragment de mile que nous ayons obtenu ne mesurait que 5 millimetres 
670 ji. C’etait une extremite posterieure. 




I 12 


posterieure est plus etroite encore, generalement recourbee sur elle-^ 
meme sans orientation uniforme de l’incurvation qui depend de la 
situation momentanee occupee par le ver dans le tissu. 

Le cloaque s’ouvre & 21/1 de Textr&nite caudale, qui k son niveau 
ne mesure plus que 27 n de large. Les papilles sous forme de tuber- 
cules microscopiques, dont les plus volumineux ne depassent la 
cuticule que de 15/*, paraissent au nombre de six paires dont quatre 
preanales, une para-anale et une postanale.* Les deux spicules 
sont tres inegaux. L*un court et large, a bords epais, concave sur 
sa face anterieure, se termine en pointe obtuse. II engaine 
partiellement le spicule long a son point d'emergeance. II est large 
de 6 p a 8ju a sa base et long de 36/1 a 43 /*. L’autre long et mince, 
arrondi, mesure au total 242 /*, atteignant son maximum de largeur 
de 5*6/1 pres de sa base d'insertion. II presente generalement une 
partie interne proximale longue de 208// et une partie externe distalc 
recourbee vers Parriere et mesurant 34// (Planche III, fig. 3A). 

Fenielle. Longueur inconnue.t Plus large que le male, elle 
alteint jusque 104 p d’epaisseur.J 

L’anus s'ouvre a yjp ou 41/1 de Pextremite caudale, au devant 
d’une depression ovalaire qui termine le ver et qui montre une 
structure interne que P£tat de conservation de nos parasites ne nous 
a pas permis de preciser. 

L’uterus comprend deux tubes genitaux quelquefois d'in^gale 
longueur, qui s’etendent en arriere jusqu’a 493 /a de Textr&nite 
posterieure. Ils se reunissent en avant, en un ovejecteur unique. 
Celui-ci, long de 286/1 et large de 16/1, presentant au moins une 
inflexion, peut-etre poursuivi jusqu’a 52/1 en arriere de la bouche. 
II pr6sente pr^s de son extr6mit£ anterieure un epaississement 
annulaire. Je ne suis pas parvenu a situer exactement Torifice 
vulvaire qui n'est certainement pas saillant. Les tubes ovariques 

• Pour determiner definitivement le nombre exact et la situation precise de ces papilles 
l’examcn d’exemplaircs extraits de tissu frais sera necessaire. Les papilles sont microscopiques 
et sans l’avis autorise de M. L. Gedoelst nous n’aurions pas ose considerer les proeminences 
cuticulaires que nou9 avions observees comme des papilles. 

t Le plus long fragment de $ que nous ayons obtenu, mesurait 9$ millimetres £ pardr 
de I’ext remite posterieure. II representait certainement les J de Padulte car les deux tubes 
genitaux contenaient £ leurs extremes brisees dcs oeufs £ cmbryons compietement forme*. 

La longueur totale du ver fenielle ne depassera probablement pas 12 millimetres. 

1 Les mensurations out et£ faites sur des sections de vers fixes dans les tissus et montrant 
les deux tubes ovariques contenant des oeufs embryonnes. En realite les dimensions oscillent 
entre 81 /i et 104 /*. 



contiennent des oeufs embryonnes montrant des microfilaircs 
vivantes. L’une de celles-ci echapp6e de sa coque ovulaire dans le 
tube ut6rin meme, mesurait 65 p de long sur 3/u de large. La femelle 
est ovovivipare (Planche III, fig. 3B). 

Nous appellerons Filaria fertenne cette filaire entierement 
differente de Dirofilaria kuelzii , Rodenwaldt (1910), qui parasite le 
tissu conjonctif souscutan6 et intramusculaire de Cephalophus 
ntaxwelli au Cameroun, et que L. Gedoelst (1916) a retrouve au 
Congo Beige. II ne nous est pas possible de dire combien de filaires 
vivent dans une meme tumeur cheloidienne. Leur nombre parait 
etre considerable, car hors d’un fragment de tumeur ayant pres de 
1 centimetre cube de volume, nous avons isol6 cinq extremity 
post^rieures de males, et trois de femelles. 

Ainsi que je 1 ’ai dit prec6demment, les vers, du moins leurs 
extremites, se deplacent dans le tissu fibreux qu’ils habitent. Ces 
mouvements ont tres vraisemblablement pour but d'assurer les 
contacts des sexes necessaires pour la reproduction. II est probable 
aussi, que la femelle effectue sa ponte dans la lesion fibromateuse 
meme, quoique sur nos coupes nous n'ayons pas rencontr6 de micro- 
filaires en dehors des tubes ovariques. 

Un examen^japide du sang peripherique de la c6phalophe 
parasite n’a pas montre de microfilaires. Nos investigations n’ont 
pas port6 sur les ganglions inguinaux drainant les r6gions oil 
si^geaient les tumeurs ch 61 oidiennes. 

Nous ne pouvons done actuellement formuler aucune hypothese 
fondle sur le mode de transmission de ce nouveau parasite. 


BIBLIOGRAPHIE 


Gf.uof.LST, L. (1916). Notes sur la faune parasitaire du Congo Helge. Revue Zoologique 
Ajricaine , Vol. V, fasc. i. 

Rodenwaldt, E. (1910). Filaria kuelzii , n.sp. Arcb.J. Scbi£s. u. Trop. Hyg XIV, p. 529. 



Fig. i 


Fig. 2 


ii4 


EXPLANATION OF PLATES 


Plate II 
Cheloide filarientit 

Coupe perpendiculaire a la peau et montrant : a. Les 
dilatations du reseau lymphatique sous-papillaire. b. Un 
noyau inflammatoire avec diverses sections de filaires. 

(Microphotographie grossissant environ 70 fois.) 

Coupe montrant un fragment de filaire en section longi- 
tudinale, envelopp6 dans sa gangue de pus. 

(Microphotographie grossissant environ 140 fois.) 













ii6 


Plate III 


Filaria fertenue , n.sp. 

Fig. 3. A. Extrdmit6 post£rieure du male. B. Extremite anterieure 
de la femelle. 

(Microphotographie grossissant environ 100 fois.) 



Annah Trap. Med. & Parasitol., Vol. XI11 


PLATE III 




Fig- 3 


C. Tinliug < 5 -» Co. % Ltd.. Imp. 









"7 


STUDIES IN THE TREATMENT OF 

MALARIA 

XXVIII. QUITENINE HYDROCHLORIDE IN 
SIMPLE TERTIAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 

AND 

Capt. W. R. O’FARRELL, R.A.M.C. 

From the Liverpool School of Tropical Medicine 

Undertaken at the request of the War Office 

(Received for publication 10 May , 1919 ) 

Sixty grains of this preparation* were supplied to us by the 
courtesy of Professor Ramsden. 

The drug was administered by the mouth in one case (No. 1597), 
grains 20 being given on the first day, grains 30 on the second day 
and grains 10 on the third day of treatment, a total of grains 60. 
The result is shown in the chart, in which : — 

T. = simple tertian trophozoites or schizonts. 

G. = simple tertian gametes. 

Neg. = no parasites found. 

Q.T. = oral administration of quitenine. 

Q.O. = oral administration of quinine sulphate. 


# The quitenine given to the patient was prepared by Skraup’s method by oxidising 
quinine with potassium permanganate whereby its vinyl group is converted into a carboxyl 
group. 



118 


Case 1597 


ApiA 



CONCLUSION 

Quitenine hydrochloride in the doses used is of no value in the 
treatment of simple tertian malaria. 

































119 


STUDIES IN THE TREATMENT OF 
MALARIA 

XXIX. ORAL ADMINISTRATION OF LIQUOR 
ARSENICALIS MINIMS 30 DAILY FOR 16 DAYS 
WITH QUININE BIHYDROCHLORIDE GRAINS 
15 INTRAMUSCULARLY ON THE ist AND 2nd, 
8th AND 9TH, 15TH AND 1 6th DAYS, IN SIMPLE 
TERTIAN MALARIA 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 

AND 

Capt. W. R. O’FARRELL, R.A.M.C. 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
(Received for publication 12 May, 1919) 

In a previous paper (1919) we recorded the results of 
administering Liquor arsenicalis in large doses, minims 30 daily, 
over a period of eight weeks in combination with two initial intra¬ 
muscular injections of quinine bihydrochloride grains 15 on two days 
only. This treatment, which was administered in thirty-two cases, 
was followed by four relapses only (i2'5 per cent.) within an 
observation period of sixty days after cessation of treatment. 

The present observations were undertaken in order to ascertain 
whether an equally favourable result could be got with a similar 
treatment of shorter duration. 



120 


The records of the observations are given in the tables and 
charts, in which : — 

T. = simple tertian trophozoites or schizonts. 

G. = simple tertian gametes. 

Neg. = no parasites found. 

A. = oral administration of liquor arsenicalis minims 30. 

Q.M. = intramuscular injection of quinine bihydrochloride 
grains 15. 

Liquor arsenicalis minims 30 daily were given for sixteen days 
and an intramuscular injection of quinine bihydrochloride grains 15 
on the first and second, eighth and ninth, fifteenth and sixteenth 
days. 

Cases 1598—1642 

Forty-five cases (Table I) were treated in this manner. All were 
adult males, the majority of whom were infected either in Macedonia 
or in East Africa. In every instance a diagnosis of simple tertian 
malaria was made microscopically, and in all cases parasites were 
present in the blood on the day treatment commenced. Blood 
examinations were made daily. 

In four of the forty-five cases treatment was^ commenced during 
an apyrexial period. In thirty-nine of the remaining forty-one the 
temperature fell to normal within three days, and in one case in 
four days; in the remaining case (1630), owing to intercurrent 
disease, the temperature was uncontrolled until the fourteenth day. 

In forty-one cases parasites disappeared from the cutaneous 
blood in one to two days, in three cases in three days, and in one 
case in four days. 

Relapses. In nine of the forty-five cases parasitic relapses 
occurred in seventeen to fifty-six days, and febrile relapses in 
twenty-six to fifty-six days, after cessation of treatment. In 
twenty-one cases the observation period was less than sixty days 
consequently the minimum percentage of cases which relapsed (those 
actually observed) is 20 per cent., and the possible maximum (those 
actually observed + the number of cases discharged before 
completion of the sixty-day post-treatment observation period) is 
66*6 per cent. Two of the cases (1600 and 1616) which did not 
relapse within the sixty-day observation period, did so on the 
sixty-eighth and sixty-fifth days respectively. 

• This was due to the demobilization which took place subsequent to the date of the 
Armistice, u.11.18. 



121 


Table I. 

Summary of retults of oral administration of Liquor arsenicalis, minims 30 daily, for 16 days + quinine bihydrochloride, grains 15, intramuscularly, 
on the 1st and 2nd, 8th and 9th, 15th and 16th days, in simple tertian malaria. 

* E. = Egypt. E.A. — East Africa. I. = India. M. = Mesopotamia. P. = Palestine. S. = Salonika. 


uraber 

of 

case 

•Place 
of in¬ 
fection 

Interval 
(in months] 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
(in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
(in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

| Febrile 
j relapse 
(above 
ioo° F.) 

| occurred 

1 in — days 

I after 
' last dose 

' 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 
not 
relapse 

Remarks 

ly* 

M. 

29 

29 

6 

15.12.18 

1 

. 

... 

... 

5 ° 


* 59*1 

E.A. 

11 

10 

4 

15.12.18 

3 

1 

18 

... 

... 

Quinine orally, grs. 30 
on 24th day. 

i6co 

S. 


7 

6 

15.12.18 

2 

1 

... 


67 

Relapsed on 68th day. 

16c 1 

E.A. 

21 

11 

8 

17.12.18 

2 

■ 

48 

47 



\6c>2 

S. 

30 

4 

4 

17.12.18 

Apyrexia 

1 

... 


46 


i6r,3 

s - 

iS 

4 

4 

18.12.18 

3 

2 

... 

... 



16:4 

j E * 

2 

8 

2 

18.12.18 

3 

2 



6 5 


1605 

1 S. 

16 

5 

4 

18.12.18 

1 

2 

... 


98 


1606 

i S - 

*4 


1 

20.12.18 

Same day 

1 


... 

61 


: 6 zj 

s. 

3 1 

2 i 

1 

2 

21.12.18 

Apyrexia 

1 



70 


16 

| s. 

35 

9 1 

7 

21.12.18 

1 

3 

... 


70 



s. 

1 

1 

1 

1 

24.12.18 

1 

4 

... 


74 


tfil 2 

s. 

41 

18 

4 

24.12.18 

Same day 

1 

3 * 

32 

... 


1*1 I 

^ s. 

14 

4 

4 

25.12.18 

1 

1 

... 


50 


1612 

1 E. 

28 

5 j 

5 

28.12.18 

1 

1 

... 


89 


*'3 

s. 

1 

28 

3 

3 

15.1.19 

1 

1 



76 


r6i4 

1 s. 

*5 

1 

4 

4 

18.1.19 

1 

1 



63 


1615 

*•* 

... 

1 

l 

... 

21.1.19 

2 

2 

56 

56 



616 

... 




23.1.19 

1 

2 

! 


64 

Relapsed on 65th day. 

617 

; 

... 

! 

... 

28.1.19 

1 

2 

... 


5 ° 


618 

p. 

4 

« i 

1 

3 , * , * , 9 

2 

2 

... 


62 


& 19 

E. 

5 

i 

2 

4 - 2*9 

3 

1 

i 


70 









122 


Table I —continued 

Summary of results of oral administration of Liquor arsenicalis, minims 30 daily, for 16 days + quinine bihydrochloride, grains 15, intrammcub 
* on the lit and 2nd, 8th and 9th, 15th and 16th days, in simple tertian malaria. 

• E. = Egypt. E.A. = East Africa. I. = India. M. = Mesopotamia. P. = Palestine. S. = Salonika. 


Number 

of 

case 

•Place 
of in¬ 
fection 

Interval 
[in months) 
between 
first 

admission 
to a 

hospital 

with 

malaria 

and 

present 

treatment 

Interval 
[in months) 
between 
leaving 
infected 
area and 
present 
treatment 

Interval 
in months) 
between 
arrival in 
England 
and 
present 
treatment 

Date 
of end 
of 

treatment 

Tempera¬ 
ture fell 
to normal 
in — days 
after 
first dose 

Parasites 

disappeared 

from 

cutaneous 
blood 
in — days 
after 
first dose 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

Febrile 
relapse 
(above 
ioo° F.) 
occurred 
in — days 
after 
last dose 

Observa¬ 
tion 
period 
(in days) 
in cases 
which did 
not 

relapse 

Remarks 

1620 

E.A. 

11 

8 

2 

4.2.19 

2 

3 

28 

... 


Quinine orally, t 









! 



on 29th day. 

1621 

E.A. 

11 

2 

2 

5.2.19 

Same day 

2 

1 

, 

70 


1622 

S. 

20 

2 

2 

5.2.19 

Apyrexia 

I 

#M ! 

... 

5* 

102-8° F. on 1 












and 102*4° T 












28th days. 

1623 

P. 

3 

1 

1 

I 3- 1 - , 9 

Same day 

I 

34 

35 



1624 

S. 

5 

2 

> 

14.2.19 

1 

I 

- 


>4 


1625 

I. 

11 

3 

3 

19.2.19 

Apyrexia 

1 

— 

... 

5 2 


1626 

... 




22.2.19 

1 

2 

*7 



Quinine orallv. 












on 20th day. 

1627 

E.A. 

20 

*7 

9 

23.2.19 

2 

I 


... 

5 


1628 

... 



... 

25.2.19 

1 


... 


3 2 


1629 

... 

”* 

... 


25.2.19 

2 

I 

... 

... 

3 2 


1630 

... 

... 

... 

... 

25.2.19 

H 

2 

... 


21 


1631 

P. 

8 

... 

2 

25.2.19 

2 

I 

! 


5 ° 


1632 

E. 

3 

3 

0 

26.2.19 

I 

I 

2 5 

26 

... 

101-4° F. and 












on 3rd and. 

•633 

E.A. 

11 

6 

0 

27.2.! 9 

3 

2 

2 3 

*4 



>634 

... 


... 

... 

27.2.19 

1 

2 

... 

... 

30 


>635 

... 


... 


27.2.19 

2 

• 


... 

■3 

102° F. on 71 

1636 

... 


... 


28.2.19 

2 

1 

I 

... 

18 


1637 

... 


... 


2.3.19 

1 

I 

... 


16 

1 


1638 


... 

... 

... 

2.3.19 

4 

3 

| 


«5 

1 

'639 

1 


1 


2.3.19 

Same day 

2 



17 


1640 

... 

— 

1 

... 

5 - 3*9 

Same day 

2 



*3 


1641 

s. 

8 

5 

2 

6.3.19 

1 

1 

... 

... 

60 


1642 

E. 

21 

... 

1 

*3-3*9 

2 

2 

... 


21 










123 


Table II. 

Summary of results of intramuscular injections of quinine bi hydro chloride grains 15 on each of 
two consecutive days only in simple tertian malaria (control series). 


Number 

of 

case 

Date 
of end 
of 

treatment 

Tempera- < 
ture fell 
to normal 
in — days 
after first 
injection 

Parasites 

iisappeared 

from 

cutaneous 
blood 
in — days 
after first 
injection 

Parasitic 
relapse 
occurred 
in — days 
after 
last dose 

f 

Febrile 
relapse 
(above 1 
ioo°F.) 
occurred | 
in — days 
after 
last dose 

1 

Observa¬ 
tion 
period 
(in days) 
in cases ; 
which did j 
not ! 
relapse 1 

Remarks 

1643 

9.3.19 

1 

1 

1 

13 

,5 ; 



16+4 

22.3.19 

2 

1 

4-5 

19 

22 

... 


1645 

30.3.19 

2 

5-6 

16 

18 

... 

1 

1646 

44.19 

« 

2-12 

12 

12 J 

... 


1647 

5.4.19 

I 

2-11 

! .8 

18 

... 


1648 

6.4.19 

Apyrexia 

... 

9 

22 " ' 



1649 j 

6.4.19 

■ 

3-10 

21 

23 

... 


1650 

7.4.19 

4 

2-4 

+ 

25 



1651 

9.4.19 

1 

2 

... 

... 

20 


1652 

9.4.19 

Same day 

1-7 

! ,6 

,6 

... 

ioi°F. on 9th day. 
io 2*6°F. on 10th day. 

1653 

10.4.19 

1 

2-5 

1 10 

13 



1654 

10.4.19 

1 

2-5 

17 

18 

... 


1655 

10.4.19 

Apyrexia 

i -5 

15 



Quinine orally, grs. 45, 
on 21 st day. 

1656 

10.4.19 

2 

2*5 

i 12 

14 


1 

1657 

11.4.19 

1 

1-4 

j l 7 

18 

... 

!|; 

1658 

114.19 

Apyrexia 

i -5 

• 13 

15 

... 


1659 

114.19 

Same day 

i -4 

1 

... 

24 


1660 

i 13419 

Apyrexia 

i -3 

14 


... 

Quinine orally, grs. 45. 
on 19th day. 



124 

CONTROL SERIES (Cases 1643-1660) 

Quinine bihydrochloride grains 15 on each of two consecutive 
days only. 

In four of the eighteen cases (Table II) treatment commenced 
during an apyrexial period; in thirteen cases the temperature fell to 
normal in two days, and in the remaining case in four days. Blood 
examinations were too infrequent to give a figure for the time of 
disappearance of parasites from the cutaneous blood. 

Relapses. In sixteen of the eighteen cases parasitic relapses 
occurred in four to twenty-one days, and febrile relapses in twelve 
to twenty-five days, after cessation of treatment. In the two cases 
(1651 and 1659) which did not relapse the observation period was 
less than sixty days, viz., twenty and twenty-four days respectively. 
Consequently the minimum percentage of cases which relapsed (those 
actually observed) is 88'8 per cent., and the possible maximum (those 
actually observed + the number discharged before the completion of 
the sixty-day post-treatment observation periods is 100 per cent. 


CONCLUSION 

It is not possible to form a definite estimate of the value of this 
treatment, as almost half the cases were insufficiently observed after 
cessation of treatment. 


REFERENCE 


Stephens, J. W. W., Yorki, W., Blacklock, B., Macfie, J. W. S., Cooper, C. F., and 
Carter, H. F. (1919.) Ann, . Trop . Med. & Parasitol . , Vol. XII, p. 371. 



125 


STUDIES IN THE TREATMENT OF 

MALARIA 

XXX. AT WHAT TIME AFTER CESSATION 
OF QUININE TREATMENT DO RELAPSES 
OCCUR IN SIMPLE TERTIAN MALARIA ? 

(SECOND COMMUNICATION) 

BY 

Lieut.-Col. J. W. W. STEPHENS, R.A.M.C. 

W. YORKE 
B. BLACKLOCK 
J. W. S. MACFIE 

AND 

Capt. W. R. O’FARRELL, R.A.M.C. 

From the Liverpool School of Tropical Medicine 
Undertaken at the request of the War Office 
(.Received for publication 31 May, 1919) 

In a previous paper (1918) we have recorded the time incidence 
of the relapses (270) among 405 cases of simple tertian malaria. We 
have now at our disposal the data with regard to an additional 
395 cases, and propose to record the time incidence of all the relapses 
(582) in the 800 cases. Before doing so we desire to indicate certain 
points which bear on the question at issue. 

1. So far as possible all cases were observed for at least sixty 
days after the cessation of treatment. As only forty-six cases were 
discharged before this time we can determine with considerable 
accuracy the incidence of relapses up to the end of this period. 
Many cases were observed for varying periods exceeding sixty days, 



126 


but as the number of cases diminished through discharges from 
hospital the figures for periods after sixty days become less accurate. 

2. All observations were made under the conditions of life 
prevailing in a military hospital in this country. We are, therefore, 
not in a position to say whether the incidence of relapses would be the 
same where other conditions may prevail. 

3. By the word ‘ relapse ’ we mean a parasitic relapse, febrile or 
non-febrile, i.e., that parasites have reappeared in the blood after a 
negative period induced by treatment. In long treatments, lasting 
say eight weeks, only those cases can be considered, in which the 
blood was negative on the last day of treatment or on the following 
day, the condition of the blood during treatment being disregarded; 
this, of course, does not apply to short treatments lasting one or two 
days, where several days may elapse before the blood becomes 
negative. 

4. All the treatments were quinine treatments. 


the tables: — 


Q.O. 

= Quinine sulphate or hydrochloride orally. 

Q.M. 

= Quinine bihydrochloride intramuscularly. 

Q.M.alk. 

= Quinine alkaloid intramuscularly. 

Q.V. 

= Quinine bihydrochloride intravenously. 

10 or 15 x 

1 = Grains io or 15 once. 

5 x 2 

= Grains 5 on each of two consecutive days. 

5 X 7 

— Grains 5 on each of seven consecutive days. 

H 

= In Table V, number of cases discharged without 


having relapsed. 

An examination of the protocols shows that in any treatment the 
majority of relapses occur during the first thirty days of the 
observation period. 

In Table I the percentage of relapses in each twenty-day period 
is given, from which it will be seen that in the different treatments 
the percentages vary from 46’2 to 100 in the first, o to 30 8 in the 
second, o to 7 "j in the third, and o to 1*5 in the fourth twenty-day 
period. A careful examination of these variations, and of the varia¬ 
tions in the treatments, e.g., (1) dose, (2) mode of treatment, 
(3) duration of treatment, has failed to reveal any correlation between 
the two. We conclude, therefore, that the variations in the incidence 
of relapses are not dependent on differences of treatment, but are due 
simply to the fact that the numbers in certain of the treatments are 



127 


Table*!. 


Showing the percentage of relapses occurring during each 20-day period after various 
quinine treatments in simple tertian malaria. 


Observation 
period after 
cessation of 
treatment. 

Days 

1 

2 

3 

4 

5 

6 

7 

8 

9 

0.0. 

5 xS 

0.0. 

10x2 

0.0. 

15x2 

0.0. 

30 x 2 

0.0. 

45 x 2 

0.0. 

00x2 

0.0. 

00x2 

0.0. 

120x2 

0.0. 

5x7 

for 8 
weeks 

1-20 

100*0 

100*0 

93 *° 

100*0 

77*8 

85-7 

75*2 

88*9 

88*6 

21-40 

... 


7*0 


22*2 

H ’3 

21*2 

1 i*i 

8*6 

41-60 


... 





3*5 


2*8 

61-80 



... 



... 

... 

... 

... 

81-100 

... 





... 

... 

... 

... 


10 

11 

12 

13 


IS 

16 

17 

18 

Observation 
period after 
cessation of 
treatment. 

Days 

Q.O. 

15 x 7 

for 8 
weeks 

Q.O. 

20x7 

for 14 
weeks 

Q.O. 

30 x 7 

for 8 

weeks 

Q.O. 

30x7 

for 3 
weeks, 
45 x 7 
for 1 
week 

0.0. 

45 x 7 

for 3-8 
weeks 

0.0. 

10x2 

for 8 
weeks 

Q.O. 

15x2 

for 8 
weeks 

0.0. 

45x2 

for 8 
weeks 

Q.O. 

90 x 2 

for 3 

weeks 

1-20 

92|3 

I 00*0 

70*6 

88*2 

7 I# 4 

1 

100*0 

80*4 

90*6 

46*2 

21-40 

3-8 

... 

26*4 

5-9 

28*6 


13*0 

6*2 

30*8 

41-60 

3-8 


3*0 

5*9 


... 

6*6 

*•5 

7*7 

61-80 

... 




... 

... 1 

... 

»*5 

0*0 

81-100 






I 


. 

0*0 


19 

20 

21 1 

22 

23 

i 

* 



Observation 
period after 
cessation of 
treatment. 

Days 

Q.M. 

16x2 

Q.M. 

alk. 

15-60 

x2 

Q.M. 

and 

QO 

30 

for 12 
days 

Q.V. 

10 or 
15 x 1 

Q.V. 

10 or 

15 x 6 | 


Range 


1 -zo 

89*3 

71*0 

| 

69*2 

100*0 

100*0 | 

I 

i 

46*2 — 

100*0 


21-40 

10-7 

29*0 

23*1 

... 1 

... 


0*0 — 

30*8 


41-60 



7*7 



! 

0*0 - 

7.7 


61-80 

... 

... 

... 




0*0 - 

i *5 


81-100 

... 

| 

I 

i 



0*0 — 

0*0 





















128 


small. It is, therefore, justifiable to consider the time incidence of 
the relapses (582) as a whole. 

The time incidence of relapses can be considered in three ways. 

1. In reference to the relapses themselves , i.e. the percentage of 

the total relapses which occur during each period of time. 

This is shown in Table II and Graph I, from which it follows that 
if in any treatment we know the number of relapses which have 
occurred in the .first twenty days after cessation of treatment, then 
we can predict the total number of relapses in the second and third 
twenty days; e.g. if eighty-three relapses have occurred in the first 
twenty days then only thirteen or fourteen will occur in the second 
twenty days, and two or three in the third twenty days, and none or 
one in the fourth twenty days. 

2. In reference to the total cases treated , i.e. the percentage of the 

total cases treated which relapse during each period of time. 

This is shown in Table III and Graph 2, from which it follows 
that if in any treatment we know the percentage of cases treated 
which relapse in the first twenty days after the cessation of treatment, 
then we can predict the incidence of relapses in the second and third 
twenty days, e.g., if 60 per cent, of the total cases treated relapse 
in the first twenty days, then about 10 per cent, of the total cases 
treated will relapse in the second twenty days, about 2. per cent, of 
the total cases treated in the third twenty days, and about 
0’2 per cent, in the fourth twenty days. 

3. In reference to remainders , i.e. the incidence among the cases 

treated less those who have previously relapsed. 

This is shown in Table IV and Graph 3, from which it follows 
that if in any treatment we know the percentage of cases treated 
which relapse in the first twenty days, then we can predict the 
percentage of the remainder that will relapse in the second, third 
and fourth twenty days, e.g., if 60 per cent, relapse in the first 
twenty days then about 26 per cent, of the remainder will relapse 
in the second, 7 to 8 per cent, of the remainder in the third, and 
about 1 per cent, of the remainder in the fourth twenty-day period. 



129 


Table II. 

Showing the percentage of total relapses (582) which occurred 
during each 20-day period. 


Observation period 
after cessation 
of treatment. 

Days 

Relapses 

Percentage 

1-20 

484 

8320 

21-40 

80 

*374 

41-60 

*5 


61-80 

1 

017 

8i-iod 

0 

0-00 

101-120 

1 

0-17 

121-140 

1 

0-17 

Total 

$8 2 

100-03 


Table Ilf. 


Showing the percentage of cases treated which relapsed during each 20-day period after 

cessation of treatment. 


Observation period 
after cessation 
of treatment. 

Days 

Original number 
of cases treated 
less those 

discharged without 
having relapsed : 
average for each 
20-day period 

Relapses 

Percentage 

1-20 

7987 

484 

6o-6 

21-40 

7866 

80 

10*2 

41-60 

763-5 

*5 

2*0 

61-80 

666.1 

1 

0-2 

81-100 

6057 

0 

0*0 










130 


Table IV. 


Showing what percentage of cases, not having previously relapsed, did so during each 
20-day period after cessation of treatment* 


Observation period 
after cessation 
of treatment. 

Days 

Average number of 
cases in hospital 
during each 20-day 
period, less the 
relapses during 
the previous 
20-day period* 

-j- 

Relapses 

Percentage 

1-20 

7987 

484 

6o*6 

21-40 

3026 

80 

264 

41-60 

1995 

15 

7*5 

61-80 

87-2 

1 

11 

81-100 

257 

0 

0*0 


REFERENCE 

Stephens, J. W. W., Yorke, W., Blacklo^k, B., Macfie, J. W. S,, Cooper, C. F., and 
Carter, H. F. (1918). Ann, Trop, Med, & Parasitol ., Vol., XI, p. 425. 




2 


792 


9 

3 V - 


25 









Percentage*. 


Graph i. * 


Graph 3. 


131 

Graph 2. 

Percentage of total relapses Percentage of cases treated Percentage of cases treated 

in each 20-day period. which relapse in not having previously 

each 20-day period. relapsed which do so in 

each 20-day period. 



Days after cessation of treatment. 




IJ 3 


THE PHAGOCYTOSIS OF ERYTHRO¬ 
CYTES BY AN AMOEBA OF THE 
UMAX TYPE 


BY 

WARRINGTON YORKE 

AND 

J. W. S. MACFIE 
(Received for publication 21 March , 1919) 

During the past ten or twelve years much has been written on 
the question of the differentiation of the vegetative stages of 
Entamoeba histolytica and Entamoeba coli. One of the main 
characters which has been held to be of use for this purpose concerns 
the ability of the parasites to ingest red corpuscles, but even on this 
point there is considerable difference of opinion. 

Amongst the rules laid down by Wenyon and O’Connor 
(1917) as a guide to the diagnosis of amoebae in the stools is the 
following :— 1 If amoebae containing red blood corpuscles are present 
in a stool, whether evidently dysenteric or not, they are E. histolytica , 
and mean that some active dysenteric process is going on.’ On 
the other hand James (1914), discussing this question, states that 
occasionally he has found one or two red cells in E. coli. In order 
to test the powers of E. coli to ingest red blood corpuscles, Wenyon 
and O’Connor mixed a portion of perfectly fresh stool, containing 
free E. coli in large numbers, with a quantity of finger blood; the 
mixture was placed at once into the incubator and was examined 
from time to time. Although the amoebae were seen to be moving 
freely amongst the blood cells, none of the latter were ingested. 
From two experiments of this nature Wenyon and O’Connor 
conclude that E. coli does not readily ingest red blood cells under 
the conditions of the experiment. The authors also state that they 
failed to induce E. histolytica to take up red cells under similar 
conditions. From these observations they infer ‘that if amoebae 
are found with phagocyted red blood corpuscles they are certainly 
E. histolytica , and are taking part in some active dysenteric process.’ 



*34 


The very fact that in the above experiments E. histolytica failed 
to ingest red cells suggests to our minds that the conditions of 
experiment were such that no definite conclusion can be drawn 
regarding the capacity of E. coli to phagocytose erythrocytes. The 
following observations made by us are of interest in so far as they 
prove that the capacity to ingest red blood corpuscles is not peculiar 
to E. histolytica. Our experiments were conducted with an amoeba 
of the Limax type obtained from the faeces of a native at Freetown, 
Sierra Leone, in 1914, and subsequently maintained in the Liverpool 
laboratory by cultivation on Musgrave and Clegg's medium. It 
was found that if a small portion of a fresh culture containing 
abundant actively motile amoebae were mixed with a little blood 
and placed in the incubator, phagocytosis of the red corpuscles did 
not take place. When, however, sub-cultures of the amoebae were 
made on Musgrave and Clegg's medium containing about one-third 
of its volume of blood, it was found that, after a few hours' 
incubation, large numbers of the amoebae contained red cells, 
sometimes as many as twenty in a single amoeba. 






Fin. Showing phagocytosis of erythrocytes by an Amoeba of the Limax type. X 1000. 


These two experiments show that although no phagocytosis of 
erythrocytes is obtained by merely incubating a mixture consisting 
of a scraping of an amoeba culture and blood, yet under conditions 
more favourable for the growth and metabolism of the amoeba active 





135 


phagocytosis does occur. Wenyon and O’Connor have themselves 
suggested a similar explanation to account for their failure to induce 
E. histolytica to ingest red cells in experiments outside the body, 
and conclude that the amoebae found in the stool to contain red cells 
‘must be amoebae which have escaped from some definite active 
lesion of the gut, where they have been living as tissue parasites.’ 
It appears to us that there is no reason to doubt that if E. coli 
happens to encounter blood when in a state of full activity, 
phagocytosis of erythrocytes may occur. In this connection it may 
be recalled that many observers have drawn attention to the 
fact that E. coli can ingest cysts of E. histolytica and intestinal 
flagellates. If this be so, it is difficult to believe that they are 
incapable of ingesting erythrocytes. 


REFERENCES 

James, W. (19U). Ann. Trap. Med. and Parasitol ., Vol. VIII., No. 2, p. 133. 

Wenyon and O'Connor (1917). Journal Roy, Army Med. Corps , Vol. XXVIII., No. 2. p. 151. 




137 


STRONGTLIDAE IN HORSES 

VIII. SPECIES FOUND IN AMERICAN HORSES 

BY 

WARRINGTON YORKE 

AND 

J. W. S. MACFIE 
(Received for publication 7 lune> 1919) 

Plates IV-VI 

In the course of this investigation parasitic worms were examined 
from fourteen horses which had recently come from the United States 
of America and had died shortly after their arrival in this country, 
and from one foal (No. 14) which, although bred in this country, 
had been kept in a field infected by the American horses. We were 
ourselves present at the post-mortem examinations of five of the 
animals, and from the others material of one sort or another was 
sent to us by Col. Brittlebank, A.V.S., C.M.G., to whom we wish 
once more to express our thanks for the facilities he so kindly afforded 
us in our investigations and for the materials with which he was 
good enough to supply us. 

We have also examined a small collection of materials from 
mules in the Argentine, and some parasitic worms from two U.S.A. 
horses which had died in France. 

In seven previous papers (1918-19) we have described in detail 
a number of the worms, particularly those new to science; and in the 
present article we propose to give a list of all the Strongylidae found 
and to discuss briefly the pathological findings. 

I. The Species of Strongylidae identified. 

In Table I a list is given of the species of Strongylidae found in 
the American horses (including the foal); the Table shows also what 
species were found in each animal examined. It is quite likely that 
the horses harboured additional species because the materials 
obtained were sometimes small in amount or of a selected kind 
(e.g. aneurysms), and it has not yet been possible to examine 
completely the very large number of specimens collected on other 
occasions. In dealing with members of this family, it would be 
necessary to examine with a microscope every single worm collected 
in order to be certain that no species had been overlooked. 



*38 


Tabu I. 

Specie* of Strongylidae found in fifteen U.S.A. horse* examined in England. 


Species 


U.S.A. herses examined in England 



i 

2 

3 

4 

5 

6 

i 

7 

8 

9 

10 


12 

i' 3 

«4 

*5 

Strongylus 

S. edentatus . 

4 - 


4 - 

1 

1 

I 

+ 

i 

1 

i ”■ 

l 

1 

i 

+ 

| 


i 

i 

I 

... 


i 

4 - 

4 - 


5 . equinus . 


+ 

+ 

+ 

+ 

i 

+ 

4 - 

, 4 - 

4 - 

4 - 

... 

4 - 

+ 

... 

4 - 

S. vulgaris . 



4 - 



4 - 


1 

4 - 

4 * 

4 - 

j 4 - 


4 - 

+ 

4 - 

Triodontopborus 

T. intermedius . 




+ 

-f 






l 



4 - 


7 . brevicauda . 

... 

... 

... 



... 

... 

... 


... 

i ... 

... 

... 

4 - 

... 

7 . tenvicollis . 

... 

... 


... 

... 

... 

... 


... 

... 

... 

... 

... 

4 - 

... 

Gyaloeepbalus 

G. capital us 



4 - 


4 - 

4 - 








4 - 


G. equi . 




... 

... 

4 - 


... 

... 

... 

... 

... 


4 - 

.... 

Cylicostotnum 

C. bicoronatum 



i 

! 

... 







! •" 



4 - 


C. calicctum . 

4 - 


! 

... 

... 


... 

... 

+ 

4 - 


4 - 

+ 

... 

4 - 

C. cor ova turn . 

+ 


i 

+ ! 

+ 



... 


+ 

4 - 

1... 

i 

... 

+ 

'... 

+ 

C. insigtte ... 

... 


... 1 


4 - 



... 

... 


... 


... 

4 - 

••• 

C. labiatum . 

... 


+ 

... 

4 - 


... 




i — 



... 

... 

C. labratum . 



+ 


4 - 


... 




... 



i ... 

... 

C. longibursatum . 

+ 

+ 

+ 

... 

+ 

... 

... 

... 


... 


... 


... 

+ 

C. minutum . 

... 

... 

... 

... 

+ 

+ 

... 

... 


... 

... 

+ 


- 

... 

C. nassatum tar. parvum ... 


... 

+ 

+ 

+ 

+ 

... 

... 



... 

4 - 


... 

... 

C . pa ter a turn . 

... 

... 

... 

+ 

+ 

... 

... 

... 

... 


... 

... 


••• 

... 

C. pocvlatum . 

... 

... 

... 

... 

... 

... 

... 

... 



... 

... 


4 - 

... 

C. pseudo-catinatum 


... 

4 - 

+ 

+ 

4 - 


... 

1 

... 

... 

4 - 

... 

4 - 

... 


All the Strongylidae found by us belonged to the sub-family 
Slrongylinae, that is Strongylidae having a chitinous buccal capsule. 

For purposes of comparison, we give in Table II a complete list 
of all the Slrongylinae in horses, donkeys and mules recorded from 
different parts of the world. 









139 


Table II. 

Species of Strongylinae found in horses and donkeys in various parts of the world. 


Species 

Egypt 

(L00S8) 

Canada 

(Ransom) 

England 

(Leiper 

Boulenger) 

U.S.A. 
horses in 
England 
(Yorke and 
Macfie) 

Strongylus 





S. edentatus , M idler, 1784 . 

+ 

+ 

+ 

4 - 

S. equtnus ,, (Looss), 1900. . 

-f 

+ 

+ 

4 - 

$. vulgaris , (Looss), 1900. 

+ 

+ 

+ 

4 - 

Triodontopborus 





T. brevicauda , Boulcnger, 1916. 


+ 

+ 

4 - 

T. intermedins , Sweet, 1908 . 

T. minor y Looss, 1900 . 


+ 

+ 

4 - 

+ 


... 

... 

T. terrains , Looss, 1900. 

+ 

... 

+ 

... 

T. tenuicollisy Boulcnger, 1916 . 


+ 

4 - 

4 - 

Gyalocepbalus 





G. capitatuSy Looss, 1900. 

+ 

+ 

4 - 

4 - 

G. equiy Yorke and Macfie, 1918 . 

... 



4 - 

Cylicostomum 





C. alveatumy Looss, 1900. 

+ 




C. auricula turn, Looss, 1900 . 

+ 



... 

C. bicoronatumy Looss, 1900 . 

+ 

+ 

4 - 

+ 

C. calicatum, Looss, 1900 . 

+ 

+ 

+ 

+ 

C. catinatum, Looss, 1900 . 

+ 

+ 



C. coronatumy Looss, 1900 . 

C. elongatum , Looss, 1900. . 

+ 

+ 

4 - 

4 - 

+ 

+ 


... 

C. euproctuSy Boulenger, 1917 . 

C. goldiy Boulenger, 1917 . 

... 


+ 


... 

+ 

4 - 

... 

C. tnsigne, Boulenger, 1917 . 

... 

+ 

+ 

4 - 

C. labtatumy Looss, 1901. 

+ 

+ 


4 - 

C. labratumy Looss, 1900. 

C. longibursatumy Yorke and Macfie, 1918 

+ 

+ 

... 

+ 



... 

4 - 

C. nuttamiy Leiper, 1913. . 

... 

... 

4 - 


C. minutuMy Yorke and Macfie, 1918 . 

... 

... 


+ 

C. nassatumy Looss, 1900 ... . 

+ 

+ 

4 - 


C. nassatum var. parvum , Yorke and Macfie, 1918 

+ 

... 


+ 

C. pateratumy Yorke and Macfie, 1919. 

... 



+ 

C. poculatum , Looss, 1900 . 

C. pseudo-catinatumy Yorke and Macfie, 1919 ... 

+ 

+ 

4 - 

+ 

... 

.... 


+ 

C. radiatunty Looss, 1900. 

+ 

+ 



C. tetracantbumy (Mehlis), 1831. 

+ 

... 

... 


Oesopbagpdont us 





O. robustuSy (Giles), 1892. 

... 

+ 

4 - 

... 


In the materials from Argentine mules Slrongylus edentatus, 
S. equtnus, S. vulgaris and Cylicostomum insigne were found. The 
parasites from two horses which had died in France were in the one 














140 


case Cylicostomum insigtie, and in the other Strongylus edentatus , 
5 . equinus, S. vulgaris , Gyalocephalus capitalus and Cylicostomum 
longibursatum. 


PATHOLOGY 


Intestinal Wall. The caecum in a number of the infected animals 
exhibited marked pathological changes. Areas of the caecal wall 
were greatly thickened and intensely engorged with blood. The 
mucous membrane was studded with small circular lesions, the 
points of attachment of worms belonging to the GENUS Strongylus 
(fig. 1). A greater or smaller number of nodules, varying in size 
from that of a pea to less than that of a pin’s head, were scattered 
throughout the mucous membrane of the caecum and upper portion 
of the colon (fig. 2). Some of the nodules were calcified ; on opening 
others with a needle they were found to be cysts lying in the mucous 
membrane, or sub-mucous tissue, and containing worms which varied 
in size with the dimensions of the cysts (fig. 3). Sections through 
the affected areas showed congestion of the blood vessels, great 
thickening of the gut wall, the presence of worms lying coiled up in 
the cysts in the mucous membrane (fig. 4) and sub-mucous tissue 
(fig. 5), and finally a general and marked endarteritis (fig. 6). 

A considerable number of worms removed from these cysts were 
examined. Without exception they proved to be larvae of the 
Genus Cylicostomum. This was ascertained to be the case because 
we were fortunate enough to find a few of the larvae undergoing 
their final moult, and were thus enabled to distinguish the 
characters of the mouth parts of both the last larval stage and the 
young adult. The larvae found in the different cysts varied greatly 
in size, some were small (4 mm.) whilst others were much larger 
(11'5 mm.). As practically all the specimens examined by us were 
in the same stage of development—the final stage with provisional 
buccal capsule—it is clear that the variation in size was not due to 
age, but to the fact that the larvae represented different species. 

It is interesting to note that larvae similar to those removed from 
the cysts were also found free in the lumen of the caecum and colon. 



A few exceptionally large cysts were found in the duodenum and 
colon of one animal (Foal 14). On opening these they were found 
to contain blood-stained pus-like matter, and larvae, or young adults, 
of Strongylus edentatus and 5 . vulgaris. 

Aneurysms of the Mesenteric Artery. Besides the general 
endarteritis observed in sections of the gut wall, aneurysms varying 
considerably in size, were found in the mesenteric artery and 
its branches in many of the horses. In every aneurysm examined 
by us, worms were found embedded in the fibrinous clot (figs. 7 
and 8). These on microscopic examination proved to be larvae of 
the GENUS Strongylus. A considerable number of these larvae were 
measured, they varied in length from 10 mm. to 16 mm. Many 
were found undergoing their final moult, and we were thus enabled 
to determine the character of the mouth parts of the last 
larval stage and those of the young adult in the same individual. 
The mouth capsule of these moulting specimens was invariably that 
of Strongylus ’ vulgaris. Judging from the appearance of the 
provisional buccal capsule, and from the slight variations in size, 
we are inclined to believe that the larvae of this species of Strongylus 
only were present in the aneurysms. 

Peri-renal Cysts. In the peri-renal tissue of one animal (Foal 14), 
a large number of cysts were found. The capsules of the kidneys 
were adherent, and in the cortex of the organs themselves there were 
some caseous cysts. Embedded in the fatty tissue surrounding the 
kidneys were a large number of worms, each coiled up in a cyst 
which contained also pus and sometimes a little discoloured blood. 
The worms were of various sizes, the largest being unmistakably 
immature adults of Strongylus edentatus. The mouth parts of the 
smaller individuals were all alike, and as one or two were moulting 
and showed both the larval and the adult forms of the buccal 
capsule, it was possible to identify the other larvae as those of 
S. edentatus . 

Lungs. We were informed that at the autopsies performed on 
many of the animals which had succumbed to sclerostomiasis, lesions 
were found in the lungs. A portion of the ‘ anterior * lobe of the* lung 
of one of these animals was examined by us. The pleura was greatly 
thickened and gelatinous, and the lung tissue itself was solid in 
places. Sections showed a greatly thickened pleura and a 



142 


pneumonic condition of the lung. Bacteriological examination 
revealed nothing definite beyond the presence of a staphylococcus 
and a bacillus of the B. coli group. 

No larvae were found in the specimens examined, and 
consequently we are unable to associate this pneumonic condition 
directly with the Strongylidae infections. 


PATHOGENICITY 

Summing up our observations, we may say, as regards the GENUS 
CyUcostomum % that the adults (sexually matur^ inhabit the colon 
and caecum, and that the larval stages are found both encysted in 
the mucous membrane and sub-mucous tissue of the same parts of the 
intestine and also actually free in the lumen of the gut. 

As regards the GENUS Strongylus, the adults (sexually mature) 
inhabit the colon and caecum, attaching themselves firmly to the 
wall of the intestine. The larvae of S. vulgaris were found in 
aneurysms of the mesenteric artery in large numbers, and 
occasionally in cysts of the colon and duodenum. The larvae of 
5. edentatus were found on one occasion in the peri-renal tissue and 
in cysts of the colon and duodenum. The larvae of S. equinus were 
not identified. Many of the worms in the aneurysms and peri-renal 
cysts had reached the adult form, and it seems incredible that they 
could migrate to the gut from these situations in this stage of their 
development; they are, therefore, to be regarded as side-tracked. 

There can be no doubt that the worms, taken as a whole, produce 
serious changes in the intestinal wall and mesenteric arteries which 
finally result in death. When one attempts to assess the importance 
of the role played by the individual genera, one is faced with 
considerable difficulty owing to the rareness with which a pure 
infection with a single genus is encountered. So far as our 
observations go, we can state that the adult Strongylus produces 
lesions in the intestinal wall at the points where it attaches itself, 
and, furthermore, that the larvae of one species, 5. vulgaris , causes 
grave lesions in the mesenteric artery and its branches. Probably 
members of the GENUS Strongylus have by far the greatest patho- 



logical significance and are mainly responsible for the changes in the 
gut wall mentioned above, and for the symptoms (emaciation, 
weakness, anemia, oedema, etc.) exhibited. 

Regarding the Cylicostomes* there can be no doubt that when 
they are sufficiently numerous the encystment of their larvae in the 
gut wall is of serious moment. 


REFERENCE 

Yorke, W., and Macfif, J. W. S. (1918-19). Ann. Trop. Med . & Parasttol Vols. XI 



EXPLANATION OF PLATES 


Plate IV 

Fig. i. Photograph of caecum of horse, showing worms of the 
Genus Slrottgylus attached to the gut wall. 


Fig. 2 . Photograph of caecum of horse, showing verminous cysts. 










146 


Plate V 

Fig. 3. Micro-photograph of final larval- stage of a cylicostome 
lying encysted in mucous membrane of colon of horse, 
x 16. 

Fig. 4. Microphotograph of section through caecum showing 

cylicostome larva encysted in mucous membrane, x 16. 

Fig. 5. Microphotograph of section through caecum showing 

cylicostome larva encysted in sub-mucous tissue, x 16. 

Fig. 6. Microphotograph of section through caecum showing 

cylicostome larva encysted in the mucous membrane 
and thickening of arterial wall, x 16. 













148 


Fig. 7 

Fig. 8 


Plate VI 


Photograph of mesenteric aneurism of horse showing 
larvae of Strongylus vulgaris . 

Photograph of mesenteric aneurism of horse showing 
larvae of Strongylus vulgaris . 




ParasitoVol. XIII 


Fig. 8 


C. Ti tiling &* Co , Ltd., I m f> 









'49 


ON THE DISTRIBUTION AND DESTRUC¬ 
TION OF QUININE IN ANIMAL TISSUES 


BY 

I. J. LIPKIN 

M.D., Ch.B., M.R.C.S., L.R.C.P., D.P.H., D.T.M. 

JOHNS rON EiLLOW IN BIO-CHEMISTRY, UVIVIRSITY OF LIVERPOOL 

From the Department of Bio-chemistry, University of Liverpool 

(.A Report to the Medical Research Committee') 

{Received for publication June 20, 1919) 

INTRODUCTION 

In a recent paper by Ramsden, Lipkin, and Whitley (1918), it 
was shown that ‘ Quinine introduced into an animal in large doses 
accumulates in most of the tissues at very much higher concentrations 
than in the blood,’ and also that ' the liver of rabbits, guinea-pigs, 
oxen and sheep rapidly attacks quinine post mortem.’ 

It was shown also (p. 256) that even a concentration of l6‘6 mgm. 
of quinine per litre, of blood, a concentration so high as to be almost 
intolerable to the patient, and which had been maintained at this 
height for at least thirty-three hours, nevertheless failed to effect a 
radical cure in a case of malaria. 

The experiments recorded »in the present paper continue the 
investigation, and are concerned mainly with 

1. The distribution of ingested quinine in animal tissues. 

2. The power of various tissues to destroy quinine. 

3. -The nature of the quinine-destroying agent and the 

conditions favouring its activity. 

4. The substances resulting from such destruction of quinine. 

DISTRIBUTION OP INGESTED QUININE 

The methods used for the extraction of the quinine from tissues 
are those described by Ramsden and Lipkin in previous papers 
(1918a, 1918^). 

1. The Ammonium Sulphate Method, serviceable for most 
tissues, e.g., spleen, kidney, suprarenal, muscle, bone marrow, testis. 



I have now tested this method with salivary and thyroid glands 
and intestinal wall, and find it equally reliable. 


Tissue 

Animal 

Weight of 
tissue 

1 

Quinine 

given 

Quinine 

found 



grms. 

mgms. 

mgm. 

Thyroid . 

...! Sheep . 

10 

5 

°’5 

(Submaxillary) Salivary gland 

.. ; 

7*5 

°*5 

°*5 


Note. —(i) In the estimation of quinine in the walls of the large and small intestine it 
was found desirable to treat the ammonium sulphate filtrate with 5% of its volume of 25% 
lead acetate solution and filter before extraction with ether, thereby removing certain im¬ 
purities which themselves give Tanret turbidity ; (2) Losses of less than 5% are to be regarded 
as within the range of experimental error. 


About 5 grms. of the tissue is transferred direct from the animal 
into a weighed flask containing about io c.c. of a saturated aqueous 
solution of (NH 4 ) 2 S 0 4 + o*6 per cent. H 2 S 0 4 . The flask is 
re-weighed, then boiled for two minutes, and its contents filtered 
under pressure through a Gooch crucible or a Buchner filter. 

The residue is pulped in a mortar with glass-powder and again 
boiled up in the original flask with successive lots of acidulated 
(NH 4 ) 2 S 0 4 solution. The combined filtrates are shaken with ether 
to extract ‘ oily * matter, then alkalised strongly with ammonia and 
again shaken up with ether to extract the quinine. Evaporate each 
lot of ether as it separates in one of the tubes guaged for 
nephelometry. Dissolve the residual quinine by boiling it with a 
sufficiency of saturated ammonium sulphate and estimate nephelo- 
metrically against quinine standards. 

2. The alcohol extraction method for brain and fat. 

Weigh out about io grammes of tissue, grind up to a fine pulp 
with powdered glass, and transfer it to a flask with the aid of 
boiling absolute alcohol. 

Boil and filter into a graduated cylinder. Repeat the boiling 
and filtering with three further lots of alcohol. Note volume of 
filtrate, pour it into five times its volume of I per cent. H 2 S 0 4 , and 
shake thoroughly for five minutes. Extract the fats and lipoids by 
shaking with three successive lots of pure ether. For every ioo c.c. of 
fat-free liquid add 5 c.c. of 25 per cent. Pb. acetate solution, filter off 
aliquot portion of the whole into a stoppered cylinder. Saturate with 



(NH 4 ) 3 S 0 4 , pipette off the layer of alcohol-ether which separates, 
and again extract with ether until the extracts are colourless. 
Alkalise with NH 4 OH, extract quinine with four successive lots of 
ether, evaporating each lot as separated in one of the tubes gauged 
for nephelometry. Dissolve the residual quinine by boiling it with 
a sufficiency of known volume of saturated aqueous ammonium 
sulphate and estimate nephelometrically. 

EXPERIMENT I. Guinea-pig, 755 grammes in weight. 0*4 grammes 
of quinine dissolved in 10 c.c. of water with enough HC 1 to effect its 
solution was injected into the peritoneal cavity. The solution was 
not acid to Congo red, or methyl orange, although acid to litmus, 
and its osmotic pressure was calculated as about half that of 
0*9 per cent. NaCl. The animal died fifteen minutes after the 
injection. The peritoneal cavity, which was in parts denuded of its 
epithelial lining, contained 5 c.c. of turbid faintly yellow fluid full of 
leucocytes and masses of rolled-up epithelial cells. Each organ 
investigated was rinsed free from peritoneal fluid by salt solution and 
wiped dry before weighing. Blood was taken from the left ventricle, 
urine from the bladder. The intestines were congested, but nothing 
else abnormal was noticeable. There were no notable symptoms 
during the period between the injection and the death of the 
animal. 


Tissue 

Grammes of 
tissue taken 

Mgm. quinine 
found 

Mgm. quinine 
per 100 grammes 
of tissue 

Suprarenal. 

°’35 

4*23 

1210 0 

Peritoneal Fluid. 

2-15 

14*6 

679-0 

Kidney . 

2-2 7 

1-93 

85*0 

Spleen . 

°*75 

q 

N 

00 

00 

o* 

Liver . 

3*25 

1*21 

37*5 

Pancreas . ... . 

I >'53 

0-23 

* 5 *° 

Muscle . 

i 

2*30 

0*20 

9 *o 

Blood . 

n 5 

0*055 

4*73 

Brain . 

2*68 

0*065 

2*86 

Urine . 

2*3 c.c. 

0*06 

2*6 

per ico c.c. 
















152 


Hence in fifteen minutes 91 per cent, of the quinine had been 
absorbed. The very high quinine content of the suprarenal confirms 
the previous results of Ramsden, Lipkin and Whitley (1918), and this 
notwithstanding the probability that, owing to the short interval 
elapsing before death, quinine accumulation had not reached its 
maximum. 

Experiment II. Guinea-pig, 463 grammes in weight. 
0 4 grammes of quinine dissolved in 2*5 c.c. of water, with enough 
HC 1 to effect its solution, was injected into the gluteal muscle. The 
injected fluid was not acid to Congo red. The animal showed 
great restlessness after fifteen minutes, and was killed by a blow 
on the occiput seventy-five minutes after the injection. The site 
of the injection showed nothing remarkable, and no fluid or exudate 
could be found there. 


Tissue 


Grammes of 
tissue taken 

Mgm.^uinine 

found 

Mgm. quinine 
per 100 grammes 
ol tissue 

Suprarenal. 

... 

0*22 

0*05 

25*25 

Thyroid . 


O'10 

0*025 

25*0 

Spleen . 


°*3 

0*068 

22*9 

Bone marrow . 


0*07 

0*009 

13*0 

Small intestine (wall) 


0*30 

0*037 

12*35 

Kidney . 

. 

3’io 

0*26 

8*3 

Brain . 


2*44 

0*14 

5*9 

Large intestine (wall) 


2*02 

0*09 

4*8 

Blood . 


3-84 

0*14 

3*7 

Muscle . 


4*9 

0*14 

2*73 

Heart muscle . 


i*i 7 

0*029 

2*5 

Liver . 


1*82 

0*04 

2*5 

Bile. 

. 

2*4 c.c. 

0*006 

0*25 

per 100 c.c. 


Although the suprarenals have again the highest percentage of 
quinine, their lead is small, and the concentration is very much less 
than with intraperitoneal injections. 












*53 


Experiment III. Buck rabbit, weight 1380 grammes. 
1250 mgm. quinine dissolved in 5 c.c. of water with just enough 
HC 1 to effect its solution was injected into the gluteal muscle of 
the leg. The fluid injected was not acid to Congo red. The 
animal died in convulsions in fifteen minutes. 


Tissue 

Grammes of 
tissue taken 

Mgm. quinine 
found 

Mgm. quinine 
per 100 grammes 
0: tissue 

Suprarenal. 

... 

0*21 

0*062 

1 

29*76 

Kidney . 


4*24 

1*25 

29*5 

Large intestine (wall) 


0*32 

0*05 

> 7*4 

Small intestine (wall) 


0*48 

0*08 

16*66 

Spleen . 


0-32 

0*036 

11*16 

Bone marrow . 


o *37 

0*03 

*3 

Muscle . 


*’33 

0*102 

77 

Brain . 


2*65 

0*12 

4*54 

Liyct . 


2 ‘35 

0*09 

3*85 

Testes . 


00 

b 

0*02 

3*03 

Blood . 

. 

6*87 ! 

0*08 

«*«7 


Urine turbid with phosphates. Quinine, blood and albumen absent. 


The short time elapsing before death was probably not long 
enough to permit of the differences of distribution in the various 
tissues attaining their maximum. 












i54 


Experiment IV. Buck rabbit, weight 1410 grammes. 
0*500 mgm. quinine in 1 c.c. of water with enough HC 1 to effect its 
solution injected into the gluteal region. The animal died seventy 
minutes after the injection. 


Tissue 

Grammes of 1 
tissue taken 

' 

Mgm. quinine 
found 

Mgm. quinine 
per 100 grammes 
of tissue 

Suprarenal ... 

o-2 5 

0*07 

28*6 

Spleen . 

0*24 

0*057 - 

2**o 

Kidney . 

4*07 

o*8o 

20*0 

Lung . 

o *57 

o*o6 

11* l6 

Fat. 

0*52 

0*058 

I Ci 

Marrow 

0-645 

0 

*0 

8*6 

Muscle . 

1*20 

0*08 

7 * 0 

Heart muscle . 

123 

0*066 

5*4 

Large intestine (wall) . 

r88 

0-086 

4*6 

Liver . . 

3*35 

0-9 

2*8 

Brain . 

251 

0*067 

2*7 

Small intestine (w'all) . 

1*40 

0-037 

2*7 

Testes . 

1 36 

0*028 

2*1 

Stomach . . 

3*89 

0*047 

1*23 

Lymph gland . 

o-68 

0*006 

0*9 

Blood arterial . 

8*43 | 

0*102 

1*2 

Blood venous . 

47. | 

0-068 

r 4 6 


Venous blood was taken from the iliac vein on the side of the 
injection; arterial blood from the left ventricle. It is noteworthy 
that the venous blood coming from the side of injection is richer in 
quinine than the arterial blood from the heart. The lymph glands 
examined were from the cervical chain. The site of the injection 
showed nothing abnormal. 













*55 


EXPERIMENT V. Guinea-pig, weight 685 grammes. 400 mgm. 
quinine in 2 c.c. of solution injected into the gluteal muscle. Animal 
killed sixty minutes later. 


Tissue 

Grammes of 
tissue taken 

Mgm. quinine 
found 

Mgm. quinine 
per 100 grammes 
of tissue 

Suprarenal. 


0*25 

0*09 

38*0 

Spleen . 


0*63 

0-15 

24*0 

Small intestine (wall) 


°’45 

0*06 

14*6 

Bone marrow . 


o-68 

0*08 

12*6 

Kidney . 


2*46 

0*21 

8-8 

Large intestine (wall) 


0*64 

c*c3 

5*3 

Brain . 


*‘54 

0*12 

4*8 

Blood . 


5*85 

°*iS 

2*6 

Fat. 


i‘ 3 * 

0*03 

*‘3 

Liver . 


644 

014 

**3 

Muscle .. 


*'45 

0 

d 

cn 

On 

**3 

Lymph gland . 

. 

0*64 

O 005 

o-8 


It should be noted that (a) no haemolysis was found even in the 
animals poisoned with quinine, (£) quinine rapidly disappeared 
from the site of injection. 

The following table furnishes a conspectus of the accumulation 
observed in the different tissues: — 














156 


Intramuscular Injections 1 Intraperttomal 

Injection 


Animal 

Guinea-pig | 

Guinea-pig 

Buck rabbit 

i 

Buck rabbit 

Guinea-pig 

Mgms. Q. per loo j 

1 


1 

i 


grammes body weight 

86*4 

5*’4 

35*4 1 

i i 

90-6 j 

53 

Time after injection 

75 min. I 

6o min. 

1 ! 

70 min. 

15 min. 

15 min. 


Tissue 

Mgm. Q. 
per 100 
grammes 
of tissue 

Tissue Q. 
Blood Q. 

Mgm. Q. 
per 100 
grammes 
of tissue 

Tissue Q. 
Blood Q. 

Mgm. Q. 
per 100 
grammes 
of tissue 

Tissue Q. 
Blood Q. 

Mgm. Q. 
per 100 
grammes 
of tissue 

Tissue Q. 
Blood Q. 

Mgm. Q. j 
per 100 1 
grammes | 
of tissue | 

Tissue ( 
Blood ( 

Blood. 


37 

— 

2*6 

* — 

1*2 

— 

***77 

— 

473 

- 

Suprarenal 


25*25 

7 *o 

w 

00 

o* 

146 

286 

* 3’83 

2976 

25-8 

1210*0 

256 

Spleen 


22*9 

6*o 

24*0 

9*23 

24-0 

20*0 

iri6 

97 

38*0 | 

8 

Kidney 


8-3 

**3 

8*8 

3*4 

20*0 

16-66 

29*5 

25*5 

00 

04 

0* 

iS 

Liver . 


»*5 

©7 

*’3 

o-88 

2-8 

2*33 

3-85 

3*3 

37*5 

3 

Muscle 


273 

075 

2*3 

o*88 

7*0 

5*83 

7*7 

6-6 

9 *o 

2 

Brain. 


5*9 

i*6 


1-85 

2*7 

2*25 

4*54 

4 *o 

3*0 

06 

Large intestine (wall) 

4*8 

>’3 

5*3 

2-04 

4-6 

3*83 

1 

* 7*4 

* 4*9 



Small intestine (u 

rail) 

12-35 

3*3 

14-6 

5 * 6 

1 47 

3 * 9 * 

j 

16*6 

* 4*2 



Lymph gland 


... 


o*8 

°*3 

0*9 

075 

... 




Bone marrow 

... 

130 

3*5 

12*6 

4*8 

I 8*6 

7*16 

8*3 

7 *o 



Heart muscle ... 

... 

**5 

07 

... 

... 

! 5 ’ 4 

4-5 

... 

... 



Fat . 




2*3 

o-8 

! in 

1 

9‘*5 ! 

... 

1 



Thyroid G. ... 


25-0 

6*8 



1 

... j 

... 




Testicle 


... 




2-' 

•7 5 

3*03 

2*6 



Lung. 



... 

... 


i iri6 

93 

... 




Stomach 



... 

... 

... 

2 ’ 3 

*'9 

; 



1 

Peritoneal fluid 








1 


* 4*7 

!_ 













1 S 7 

Partition of quinine between corpuscles and plasma of the blood 

In the experiments recorded above, samples of blood were taken 
up (4 c.c. usually) in a syringe containing 1 c.c. of I per cent 
potassium oxalate. This was centrifuged at once and the quinine 
content of the plasma and corpuscles estimated separately. 

Rabbit (1) 100 grammes of Plasma contain 2*4 mgm. quinine. 

100 grammes of corpuscles contain 1*26 mgm. quinine. 

Plasma quinine _ 2 

Corpuscle quinine 1 

Rabbit (2) 100 grammes of Plasma contain o*8i mgm. quinine. 

100 grammes of corpuscles contain 0*39 mgm. quinine. 

Plasma quinine _ 21 

Corpuscle quinine 1 

Guinea-pig 100 grammes of Plasma contain 1*98 mgm. quinine. 

100 grammes of corpuscle contain 0*62 mgm. quinine. 

Plasma quinine _ 3-2 

Corpuscle quinine 1 

V 

The blood from a case of blackwater fever gave the ratio 

Plasma quinine _ 2*2 

Corpuscle quinine I 

In a previous paper (6), one observation on the blood of a guinea- 
pig gave a ratio 

Serum quinine 3 

Corpuscle quinine 1 


ACCUMULATION OF QUININE IN EXCISED TISSUES 

A. Guinea-pig's suprarenal gland cut in two and immersed in 
10 grammes of 0*9 per cent. NaCl containing 1 mgm. quinine. 

B. Guinea-pigs suprarenal cut in two, immersed in 15 grammes 
of 09 per cent. NaCl containing 15 mgm. quinine. 

C. 0*96 gramme of guinea-pig's spleen immersed in 10 grammes 
of o*9 per cent. NaCl containing 1 mgm. of quinine. All were kept 
for forty-eight hours at 35°C., and in each case the quinine content 
of the tissue and fluid were estimated separately. 



158 


i 

i 

Grammes of 
tissue taken 

l 

Grammes of 
medium 

Mgms. quinine in 

100 grammes of bathing 
fluid 

Mgms. 
quinine per 
100 grammes 
tissue 

Quinine 
concentration 
in tissue 
Quinine 
concentration 
in Medium 

1 

! . ! 

Initially 

1 

Finally 

1 

A. j 

I 

0*21 

10 

l 1 

! 10 

i 

*’5 

307*1 

1 

123*0 

| 

1 

B. 

0*28 

*5 

j loo 

18*o 

2677*4 

j 1 54*3 

c. 

0*96 

10 

10 

2*8 

77 ’° 

22*5 

I 





__ „ 

_ 

1 


It is seen that the excised tissues, and especially the suprarenal 
gland, under conditions where their vital processes must be reduced 
to a low ebb, accumulate quinine from the environment at enormous 
relative concentration just as they do during life, a fact which 
indicates that the accumulation is dependent on some specific 
chemical or physical affinity between quinine and one or other of 
the cell constituents—it is tempting, to associate it with the lipoids 
of the cortex. To get further light on the question, attempts were 
made to find out whether the accumulation was mainly in the cortex 
or in the medulla. Glands containing large amounts of quinine, 
ranging from 250 to 385 mgm. per 100 grammes of tissue in the 
corresponding glands of the other side, were incised and immersed 
in Christensen’s Herapathite reagent for eighteen hours at laboratory 
temperature. After freezing and section, neither crystalline nor 
(black) amorphous Herapathite could be seen in either cortex or 
medulla, notwithstanding the large amount of quinine present. 

DESTRUCTION OF QUININE IN TISSUES 

The material employed for most of these experiments was 
obtained in the animal experiments already described. Each tissue 
investigated was divided symmetrically into two approximately 
equal portions; the quinine of one portion was estimated at once, 
and that of the other estimated after incubation in o‘8 per cent, 
sodium fluoride solution at 35°C. As the initial concentrations of 
quinine differ considerably in the various tissues, the results are not 
strictly comparable inter se, but indicate merely whether quinine is 
destroyed or not. 



i59 



Tissue 

examined 

j Grammes 

1 tissue 
taken 

! 

Quinine in 
mgm. per ioo 
grammes of 
tissue 
estimated 
at once 

Quinine in 
mgm. per ioc 
grammes of 
tissue after 
incubation 

Hours of 
incubation 

% . 

Destruction 

Exit. I.— 

Muscle 

475 

mm 

2*00 

18 

26*7 

Guinea-pig 

S. Intestine 

0-32 


4*i 

18 

66*76 


Blood 

r *4 

1 

3-6 

18 

27 


Spleen 

o *3 

KB 

22*2 

18 

3 *° 

Exft. III.— 

L. Intestine 

271 

* 7*4 


1 

21 

83*0 

Rabbit 

S. Intestine 

°*9 

16*6 

■ 

21 

8y6 

Expt. IV.— 

Kidney 

3-98 

20*0 

9 *o 

21 

55*° 

Rabbit 

Spleen 

0*14 

24*0 

23*81 

24 

o*8 


Testis 

1*30 

2*1 

2*0 

26 

5*° 


L. Intestine ! 

2*53 

4*6 

0*0 

24 

100*0 


S. Intestine 1 

223 

27 

2*2 

24 

* 8*5 


Muscle | 

1*20 

7-0 

4-0 

24 

43 *o 


Marrow 

0*65 

8-6 

8*32 

24 

3*3 

Guinea-pig 

Kidney 

2*45 

85*0 

8o*o 

*8 i 

6*o 


Suprarenal 

0*32 

1210*0 

1200*0 

*81 

o*8 


Muscle 

3*49 

9*0 

6*2 

*7i 

3 **o 


Peritoneal F. 

2*41 

*47 

*47 

164 

0*0 


Pancreas 

**53 

15*0 

iro 

>81 

37*0 


Thyroid 

10*0 

1 mgm. 

1 mgm. 

24 

0*0 


Salivary G. 

7‘5 

1 

1 

24 

0*0 


In the following estimations, the tissues examined were pulped in 
a mortar and added to a solution of quinine in o*8 per cent. NaF 
and incubated for varying periods at 35 0 C. All were fresh, but 
taken from various guinea-pigs. 


Tissue 

Grammes 

tissue 

taken 

Quinine 
added mgm. 

i Found 

Incubation 

period 

hours 

Loss% 

Lymph gland . 

6*3 

1 

o *95 

63 

5 *o 


3*6 

1 

0*91 

*7 

9 *o 

„ 

5 *o 

1 

1*0 

23 

0*0 

Kidney. 

47 

> 

O '93 

21 

7 *o 

n 

3*6 

1 

00 

b 

21 

16*6 

Suprarenal . 

2*5 

1 

°'95 

63 

5 '° 

u 

0*76 

. j 

0*95 

23 

5 *o 

Bone marrow . 

0*84 


0*97 

21 

3 *o 


0*69 

1 

0*96 

21 

4 *o 

Spleen . 

27 

1 

0-95 

21 

5 *o 


















MAIN CONCLUSIONS CONCERNING THE ACCUMULATION AND 
DESTRUCTION OF QUININE IN TISSUES 


Tissue 

i 

| Relative accumulating power 

1 

Relative destructive power 

Liver . 

(++) 

+ + + + + 

Spleen . 

-1 + + + 

nil 

Bone marrow . 

+ + 

nil 

Kidney . 

(+++) 

+ + 

Suprarenal . 

...1 +++++ 

1 

nil 

Muscle. 

++ 

+ + 

Intestinal wall 

...! (++) 

+ + + + 

Pancreas. 

(++) 

+ + 

Lymph gland . 

... nil 

1 

nil or slight 


In column 2, tentative concluiioni regarding tiisuei with decretory or dettroying 
powert are indicated by bracketa. 


PERFUSION EXPERIMENT 

A liver from a young sheep, freshly killed, was perfused for two 
hours at 37 0 C. with 2 litres of oxygenated Locke’s solution 
containing 200 mgm. of quinine hydrochloride not acid to Congo 
red. After perfusion, the liver was gently squeezed. The total 
quantity of fluid collected was 2024 c.c. 

Weight of liver before perfusion ... 978*6 grm. 

„ ,, after „ ... 95^ 4 » 

A portion of the liver was estimated for its quinine content—the 
total calculated for the whole liver was 18*68 mgm. quinine. 

As the total perfusion fluid finally contained 36*8 mgm. of 
quinine, 72*26 per cent, of the quinine had been destroyed in two 
hours. The final concentrations of quinine in the liver and perfusion 
fluid, weight for weight, were approximately equal. 

A weighed portion of the liver was pulped and incubated in 
o*8 per cent. NaF at 35 0 C. for forty-five hours. The whole of its 





quinine was then found to have disappeared. In the perfusion fluid 
similarly incubated only 8'8 per cent, of its quinine had disappeared. 
During the perfusion 978'6 grammes of liver destroyed 144 5 2 mgm. 
of quinine, i.e. 1 gramme of liver destroyed o'148 mgm. of quinine 
(cf. Experiment A of last paper, p. 231, where 0 4 mgm. quinine 
was destroyed by I gramme of guinea-pig’s liver post-mortem). 

At this rate 17 grammes (27 grains) of quinine would have been 
destroyed in twenty-four hours. 


OBSERVATIONS ON THE QUININE DESTROYING AGENT 
FOUND IN LIVER 

Thermolability 

10 per cent, emulsions of guinea-pig liver and ox liver in 
0*9 per cent. NaCl. Boiled for three minutes on a water bath. 
Quinine added and the mixture allowed to stand for twenty hours 
at 37 0 C. 


Tissue 

Animal 

Mgm. quinine 
added 

Found 

Loss% 

Liver. 

Sheep . 

0*2 

i'i93 

3*35 


Guinea-pig 

ro 

o*93 

7 *° 

V . 

Ox . 

ro 

o*95 

5 *o 

,, 

Guinea-pig 

ro 

0*901 

9 *i 


The experiments were repeated with unboiled liver extracts. 


Animal 

! Mgm. quinine 
| added 

Found 

i 

Loss% 

Sheep . 

1 

0*2 

1 

0*14 

30 

Guinea-pig . 

ro 

0*47 

53 

Ox. 

ro 

0*74 

26 

Guinea-pig . 

ro 

0*58 

4 1 


It is evident that the active agent is thermolabile. 




\6i 


Influence of the Reaction of the Substrate 

The experiments recorded below show that the enzyme acts best 
in neutral solution, and that alkalinity is more inhibitory than 
acidity. 

To three flasks A, B and C, each containing io c.c. of a 
io per cent, emulsion of fresh guinea-pig’s liver in a 0*9 per cent. 
NaCl and 1 mgm. of quinine, was added : — 

(a) 10 c.c. of 0*9 % NaCl. 

(b ) 10 c.c. of o*4 HC 1 . 

(c) 10 c.c. of 0-9 % Na*CO a . 

All were kept at 35° C. for twenty hours and the quinine then 
estimated. 

Mgm. Quinine 


* 

* 

added 

Found 

Loss % 

A. 

Neutral . 

. 1 

071 

28*6 

B. 

Acid. 

. 1 

0*83 

167 

C. 

Alkaline . 

. 1 

091 

91 


The experiment was 

repeated with ox liver : 

: — 




Mgm. Quinine 





added 

Found 

Loss % 

A. 

Neutral . 

. 1 

0*62 

37*5 

B. 

Acid. 

. 1 

078 

214 

C. 

Alkaline . 

. 1 

0*89 

107 


Repeated with dried 

ox liver : — 





Mgm. Quinine 





added 

Found 

Loss % 

A. 

Neutral . 

. 1 

077 

230 

B. 

Acid. 

. 1 

0*87 

130 

C. 

Alkaline . 

. 1 

091 

91 


Repeated with guinea-pig liver: — 

A. 10 c.c. of 10 % extract + 1 mgm. Q. + 10 c.c. 9 % NaCl. 

N 

B. 10 c.c. of 10 % extract + 1 mgm. Q. + 10 c.c. — H,S 0 4 . 

10 

N 

C. 10 c.c. of 10 % extract -f- 1 mgm. Q. + 10 c.c. — NaOH. 

10 

Reaction Mgm. Quinine 




added 

Found 

Loss % 

A. 

Neutral 

. 1 

0*62 

37*5 

B. 

Acid ... . 

. 1 

°74 

267 

C. 

Alkaline . 

. 1 

0-83 

167 



163 

The Influence of Oxygen 

There was some indication in the paper by Ramsden, Lipkin and 
Whitley (1918) (p. 231), that a free supply of air promoted the 
destruction of quinine by liver pulp. 

The following experiments show that oxygen is absolutely 
essential. 10 c.c. of 50 per cent, fresh liver extract in 0*8 per cent. 
NaF solution was incubated for twenty hours at 35 0 C. In Case A, 
a current of air was gently drawn over the mixture without 
mechanical disturbance of the surface; in Case B, no air was drawn 
over the mixture: — 

Mgm. Quinine 



added 

Found 

Loss % 

A. . 

1 

o *43 

57-0 

B. . 

1 

0-62 

w 

00 

6 

This was repeated. 

A. . 

1 

040 

6o-o 

B. . 

1 

o-61 

39-0 


A. 20 c.c. of fresh guinea-pig’s liver extract in o'Q per cent. 
NaCl + 1 mgm. quinine in solution mixed, and a current of air 
drawn over the mixture at laboratory temperature for twenty-one 
hours. 

B. 20 c.c. of the same liver extract placed in a flask and I mgm. 
of quinine in solution in a tube inside the flask. The flask was 
exhausted and filled with hydrogen. The quinine was then tilted 
into the liver extract and the sealed flask allowed to stand for 
twenty-one hours. 

Mgm. Quinine 

added Found Loss % 

A. . 1 0-4 600 

B. . 1 083 167 

As complete removal of oxygen was uncertain, the experiment 
was repeated with extra precautions. 

A. 10 c.c. of a o'50 per cent, extract of guinea-pig’s liver in 
0'8 per cent. + 5 drops of ammonium bisulphide solution + 1 mgm. 
quinine in solution. The flask was exhausted and then filled with 
oxygen and sealed. 

B. 10 c.c. of 0'8 per cent. NaF solution + 5 drops of strong 
ammonium bisulphide solution 4- 1 mgm. of quinine in solution. 



164 


C. 10 c.c. of same liver extract + 5 drops of strong ammonium 
sulphide solution, 1 mgm. quinine in solution placed in a small test- 
tube inside the flask. The flask was exhausted and then filled 
with hydrogen freed from all traces of oxygen by passage through 
a red hot copper tube. This was repeated three times before the 
final sealing. 

The quinine solution was then tipped into the liver extract. All 
the flasks were allowed to stand for twenty hours at laboratory 
temperature, and were heated to ioo°C. for two minutes before they 
were opened. 

Mgm. Quinine 



added 

Found 

Loss % 

A. 

. I 

0-23 

77.0 

B. 

. l 

097 

3 -o 

C. 

. I 

o *93 

6*2 

The losses 

in B and C have no significance, as 

they are 


attributable to experimental error. 


Influence of Hydrogen Peroxide on the Ferment 


A. 10 c.c. of fresh extract of guinea-pig’s liver in 9 per cent. 
NaCl 4- 10 c.c. of distilled water and 1 mgm. quinine. 

B. 10 c.c. of same liver extract 4- 10 c.c. of neutral H 2 0 2 
4- 1 mgm. quinine. 

C. 10 c.c. of same liver extract boiled and cooled 4- 10 c.c. of 
carefully neutralised H 3 O a 4- 1 mgm. quinine. 


Mgm. Quinine 


added 

Found 

Loss % 

1 

00 

6 

412 

1 

076 

23-1 

1 

0-95 

50 


Evidently hydrogen peroxide is detrimental. In the control 
experiment C, the hydrogen peroxide had not affected the quinine. 

Repeating the experiment with bile salts instead of hydrogen 
peroxide showed that they had no effect on the destruction. 



RAPIDITY OF QUININE DESTRUCTION 


A series of flasks containing io c.c. of 25 per cent, guinea-pig 
liver extract in o - 8 per cent. NaF + 1 mgm. quinine. The quinine 
content of the flasks was estimated after known intervals of time. 


Mgm. quinine : 
added I 

Found 

Time of exposure 
quinine in the liver 
extract 

Loss% 

% of total 
destruction 

t 

1 1 

q 

00 

u» 

1 minute 

16*67 

31-56 

, 

o*68 

>5 » 

31*04 

60*62 

1 

0*62 

3 ° » 

37*5 

73*24 

1 

0*58 

60 „ 

41*18 

80*43 

1 

o *55 

3 hours 

44*4 

j 

86*72 


0*52 1 

24 »i 

47‘37 

92*54 

1 

0 

00 

48 ♦? 

5 rz 1 

100*00 


It will be seen that the rate of destruction is at first considerable 
but soon becomes slow—in the first fifteen minutes 37 per cent, 
quinine was destroyed, in the second 6‘8 per cent., and in the next 
thirty minutes only 3 7 per cent. 


PURIFICATION OF THE ENZYME 

It had already been shown that 50 per cent, alcohol extracts 
the active principle from liver. It is possible by a process of 
fractional precipitation with alcohol to obtain the ferment in purer 
form. 500 grammes of minced liver were extracted with 25 per cent, 
alcohol and filtered till clear. The filtrate was red with 
haemoglobin. Alcohol was then added until by volume 36 per cent, 
was present, and the mixture was filtered. By further successive 
additions of alcohol and filtering after each addition, solutions were 
obtained with 48-8 per cent., 59 per cent., 68'8 per cent., 83 per cent, 
and 92 per cent., respectively. 

A 20 c.c. sample of each filtrate was incubated for twenty-one 
hours with 1 mgm. quinine, and the destruction of quinine estimated 
with the following results: — 





1 66 


Percentage volume of 
alcohol in filtrate 

Mgm- quinine 
added 

Found 

LpOjS 

25’o 

1 

0*62 

37*8 

36*0 

1 

0*64 

35*5 

48*8 

1 

o-66 

33*3 

59 *o 

1 

o *77 

23-0 

68-8 

1 

o *97 

O’O 

83*0 

1 

0*96 

4 *° 

92’0 

1 

o*97 

5 *o 

92*0 

0 

! 

0*0 

1 


It is clear that either the active agent was completely precipitated 
when the alcohol reached 68‘8 per cent, and upwards by volume, and 
that up to 59 per cent, very little was precipitated, or that more than 
59 per cent, inhibits its activity. 


PREPARATION OF ENZYME BY ALCOHOL PRECIPITATION 

415 grm. of minced guinea-pig liver were extracted with 
50 per cent, alcohol and allowed to stand for two hours and then 
filtered clear. 

To the straw-coloured filtrate absolute alcohol was added until 
70 per cent, was present. The precipitate was allowed to stand for 
four hours, then filtered off, and dried in a dessicator. 

The product was a whitish-yellow powder which gave no 
purpurogallin with pyrogallic acid, but contained catalase and 
haemoglobin (guiacum reaction). 0 4 grm. of the powder was 
shaken with 20 c.c. of water and filtered—the filtrate was colourless, 
faintly opalescent, and contained protein in small amount. 10 c.c. 
of it was incubated with 1 mgm. quinine for eighteen hours— 
50 per cent, of the quinine was destroyed. 

Two weeks later the experiment was repeated : only 37'5 per cent 
of the quinine was destroyed—apparently the powder deteriorates on 
standing. An attempt was made to purify the enzyme after the 
principle found effective for fibrin ferment by Schmidt. 250 grms. 
of minced fresh guinea-pig liver were extracted with absolute alcohol 
for twenty-four hours, and then shaken with ether, dried in a 





167 


dessicator and ground to a fine powder. The powder, when shaken 
up with distilled water and filtered gave a clear solution poor in 
protein. 1 grm. of the powder was incubated at 35 0 C. with 1 mgm. 
quinine -f • 20 c.c. of distilled water for twenty-one hours. Its action 
was feeble—only 26 per cent, of the quinine was destroyed. 

1 grm. of dried crude powdered ox liver, tested similarly, 
destroyed nearly as much, namely 1.7 per cent. It was evident that 
by exposure to absolute alcohol most of the enzyme had been 
inactivated. 

THE PRODUCTS OF QUININE DESTRUCTION BY LIVER 

Having found that the liver destroyed quinine only in the 
presence of oxygen, an attempt was made to identify the products 
formed. Quitenine (C 18 H„ a N a O a COOH), a well-known oxidation 
product of quinine (C 18 H aa N a O a CH : CH a ), was an obvious 
possibility, since Kerner (1870) had described its occurrence in the 
urine of patients taking quinine, and although this had been denied 
by Merkel (1902) and by Giemsa and Schaumann (1907), 
Dr. Nierenstein (1919) had informed me that he had found it in the 
urine in the earliest stages of quinine excretion. 

Professor Ramsden kindly prepared some pure quitenine by 
Skraup’s (1880) method, and together we studied some of its 
properties. In addition to the properties assigned to it by Skraup 
and Nierenstein, we find that although it gives with bromine water 
and ammonia an apparent Thalleioquin reaction, the initial colour 
is much bluer though it goes green on standing, and that, unlike 
the quinine pigment, this blue compound cannot be extracted by 
shaking with chloroform. This insolubility in chloroform furnishes 
a useful means of distinguishing it from quinine. 

Crystals of quitenine from alcoholic solution are transparent, and 
show varying ‘relief’ when examined through a rotating Nicol’s 
prism. When heated in water the crystals fragment, and become 
opaque at temperature well below 8o° C. Quitenine picrate is 
insoluble in ether. Tanret’s reagent is not nearly so delicate a 
precipitant for quitenine as for quinine, although, as in the case of 
quinine, the presence of much ammonium sulphate greatly enhances 
its delicacy. With Christensen’s reagent some darkening occurs, 
but no polarising crystals are obtainable. 



168 


Experiment i. i J lbs. of freshly minced sheep liver in 200 c.c. 
0 9 per cent. NaCl with 5 grms. of quinine dissolved with the aid 
of a minimum of HC 1 , was incubated at 35° C. for eighteen hours. 
The suspension was raised to ioo° C. for a .few moments, 
‘defaecated’ and filtered. The filtrate was saturated with picric 
acid and filtered clear. The precipitate was suspended in 100 c.c. 
of ro per cent. HC 1 , and the picric acid was completely extracted 
by shaking with ether. The acid aqueous layer was rendered 
alkaline with NaOH, and the quinine then removed by ether. A 
current of CO a was passed through the alkaline aqueous liquid, and 
the flocculent precipitate formed was dissolved in boiling alcohol 
water mixture (two parts of water to one of alcohol). The clear 
filtrate obtained on cooling deposited clear colourless crystals which 
gave all the reactions and possessed all the properties of quitenine. 

The anhydrous crystals melted at 233 0 C. The total amount of 
quitenine obtained weighed about 500 mgm. 

Experiment 2. 800 grms. of guinea-pig liver, finely minced, 
suspended in 200 c.c. o - 8 per cent. NaF containing 1 grm. of 
quinine in solution, incubated for eighteen hours at 35 0 C. 

Traces of quitenine were found. 

Experiment 3. 350 grms. of minced guinea-pig’s liver in 

o - 8 per cent. NaF 4- 4 grms. of dissolved quinine in it, were 
incubated for twenty hours at 35 0 C. 

Following the same procedure as in Experiment 1, no precipitate 
was obtained with the CO a . The solution was evaporated to 
dryness and the residue extracted with absolute alcohol. No 
quitenine, but an interesting and apparently new quinine derivative 
was obtained as an amorphous residue soluble in alcohol, water, 
acids and alkalies, and giving with bromine water and ammonia a 
green pigment which was not extracted by chloroform. Its picrate 
was soluble in ether. With Christensen’s Herapathite reagent on 
a slide it gave colourless long needle crystals. It gave a strong 
yellow turbidity with Tanret’s reagent. 

Experiment 4. 375 grms. of guinea-pig liver incubated for 

forty-eight hours to 35 0 C. with 2'5 grms. of quinine HC 1 —quitenine 
was found in small quantity, only 24 mgm. (anhydrous) being 
obtained. 



169 

QUITENINE IN URINE 

To test the influence of quitenine on the malarial parasite, 
a patient, aged 23, was given, on the nth of April, four 5-grain 
doses of quitenine hydrochloride in solution orally, six such doses 
on the 12th, and two 5-grain doses on the 13th. The patient had 
typical malarial rigors from the 10th to the 15th, inclusive. The 
quitenine had no effect on the clinical symptoms or on the presence 
of parasites in the blood. On the 15th, the patient was given 
45 grains of quinine—no further rigor occurred, and no malarial 
parasites could be found in the blood on the 17th. 

The urine collected in the twenty-four hours between the 12th 
and 13th measured 1500 c.c.; 750 c.c. of this was examined for 
quitenine by the method already described, by which I found I could 
easily detect as little as 3 mgms. in 100 c.c. of urine, but no 
quitenine whatever was found. No quitenine could be found in a 
later sample of urine. The faeces were not investigated, but it may 
be safely assumed, in view of Kerner’s (2) observations on animals, 
that most of the quitenine had been absorbed. It would seem that 
in the doses given, quitenine is not only itself ineffective but is broken 
up into therapeutically ineffective compounds, since little, if any, 
appeared in the urine. 

NOTE ON A CASE OF BLACKWATER FEVER 

Three 15-grain doses of quinine sulphate in solution were given 
orally to a man, aged 23, who had been suffering from malaria for 
two years, and whose blood contained malignant tertian parasites. 
Nine hours after the last dose of quinine, the patient vomited, had a 
rigor, passed dark urine and became jaundiced. 

A sample of urine passed three hours after the onset of the rigor 
was dark red in colour, and contained oxyhaemoglobin, methaemo 1 
globin, urobilin in great excess, albumen in fair amount, and debris 
consisting of cellular and amorphous matter and haematin. 

A sample of blood taken sixteen hours after the onset of black- 
water was found to contain o'67 mgm. of quinine per 100 grms. of 
blood (7' 1 mgm. quinine per litre of blood). 

Plasma quinine concentration _ 2.2 

Corpuscle quinine concentration 1 



170 


The blood was centrifuged immediately, the serum was deep 
reddish brown in colour and gave a strong spectrum of mixed 
oxyhaemoglobin and methaemoglobin. Urobilin was present. 

A sample of urine passed ten minutes after taking the blood 
sample was estimated to contain 35 mgm. of quinine per litre of 
urine. 

Urine quinine concentration _ 8.73 

Blood quinine concentration 1 

This remarkably low ratio has been noticed before in blackwater 
fever (cf. Ramsden, Lipkin and Whitley (1918), page 246). 

Haemoglobin was found in the urine until sixty hours, and 
quinine until sixty-four hours after the last dose of quinine. 


EXAMINATION OF THE FAECES 

Twenty-six grms. of moist faeces passed eighteen hours after the 
onset of blackwater was suspended in 1 per cent. H a S 0 4 and 
saturated with (NH 4 ) a S 0 4 , 50 c.c. of absolute alcohol added, and 
the whole mixture boiled and filtered. The residue was washed 
with mere hot alcohol. The combined filtrates were shaken up with 
ether. The ether-alcohol layer which separated was very dark 
brown, almost black in colour. This was pipetted off into another 
vessel and allowed to stand. A large quantity of blackish brown 
material separated out. 

A few drops of this ether extract poured on to a filter paper 
turned gradually from deep brown to green, yellow and red (a play 
of colours due apparently to oxidation by the air). 

The black precipitate from the ether was washed free from 
stercobilin with ether and dried. It was insoluble in water, acids, 
alkalies, chloroform, benzene, amyl alcohol, toluene, ether, but very 
freely soluble in alcohol. The alcoholic solution showed no spectral 
absorption bands, and was dark brown in colour. It turned purple 
on standing, especially if the alcoholic solution was acidified with 
hydrochloric acid. This purple pigment proved to be cholecyanin— 
the acid solution showing three well marked absorption bands. One 
thin band between the C and D lines, another broader band on the 
green side of the D line, and a third very broad band between 
b and F lines. 



On rendering alkaline with ammonia the purple colour was 
replaced by yellow, showing only one feeble absorption band on the 
D line; the purple colour reappeared on acidification. 

The purple pigment could be extracted in ether, chloroform, 
toluene or benzene, giving solutions nearly pure blue in colour. 

It seems highly probable that the initial black substance, certainly 
not itself cholecyanin although so easily converted into it, is a 
cholecyaninogen. None could be found in the urine. 

NOTE ON THE THALLEIOQUIN REACTION 

A great drawback to this very useful test for the quinine group 
of alkaloids is its lack of sensitiveness. 

The success of the test depends, among other factors, on— 

1. The concentration of the quinine in the solution to be 

tested. 

2. The avoiding of excess of bromine. 

Ramsden and Lipkin (1918^) found that, using 10 c.c. of 
solution, the least concentration of quinine certain to give a positive 
reaction with suitable precautions was 1 in 400,000, and that 0*25 
mgm. of quinine could thus be detected. 

By the following procedure, in which quinine is first isolated as 
described for the herapath test and then dealt with in strong 
solution, 0*004 mgm. of quinine can easily be detected in 40 c.c. of 
water (1 in 10,000,000): — 

1. Dissolve 5 grins, of (NH 4 ) 2 S 0 4 in every 10 c.c. of the 
quinine solution, alkalise with ammonia and extract the quinine by 
shaking with two successive lots of purified ether, transferring each 
lot as it separates to a small silica crucible in a water bath. 
Aspiration of air from its interior greatly accelerates drying. 

2. Dissolve the residue when quite dry in a minimum of ether 
squirted repeatedly down the side of the crucible by means of a 
small teat pipette. 

3. Bring the ether solution in minimum drops on to a warm 
microscope slide, in such a way that the residue left by its 
evaporation is spread over a minimal area. 

4. Cover the area of the residue with a very small drop of 
0*4 per cent. HC 1 , now add a small drop (1 /30C.C.) of weak bromine 



172 


water and immediately after a small drop of strong ammonia. 
A green colour is obtained at once. On addition of a drop of 
strong H a S 0 4 , the green colour turns to a distinct red. 

I have found that the following modification of the thalleioquin 
test, although its rationale is by no means clear, serves to distinguish 
quinotoxin from quinine. To 5 c.c. of the solution containing 
quinine or quinotoxin add a small drop of o‘5 per cent. Congo red 
in water. Add bromine water till the blue colour is discharged and 
replaced by a yellow colour. Add ammonia at once and extract 
the pigment with chloroform. Quinine gives a green, quinotoxin 
a red colour, both extracted by chloroform, but the red much more 
rapidly. One part of quinotoxin in twenty parts of quinine can be 
detected by this means. 


DISCUSSION 

• {Written jointly with Professor Ramsden ) 

One of the main objects of the work has been to get fresh light 
on the question, how it is that quinine, although almost always 
successful in causing the disappearance of malarial parasites from the 
circulating blood and effecting a temporary cure, nevertheless almost 
always fails to obviate eventual relapse. That this failure is not due 
to any lack of quinine in the general blood-stream, at least up to 
almost intolerable amounts throughout a period of many hours, has 
been shown by Ramsden, Lipkin and Whitley (1918^) in collabora¬ 
tion with Professors Stephens and Yorke. 

Assuming for the moment that quinine is directly parasiticidal, 
and also that its metabolites may be neglected, various explanations 
may be suggested for this failure: — 

1. The existence of quinine-resistant parasites or of a 

specially resistant phase in their life history. 

2. Parasites may find safety outside the blood, either in other 

body-fluids or inside cells free from quinine. 

3. Parasites may find ‘backwaters’ maintained free from 

quinine inside the blood vascular system. 

No conclusive evidence can be adduced for any of these 
possibilities, but the data showing the quinine-destroying powers 
and quinine content of the various tissues have obvious bearings on 



I 73 


the last two of them, although subject to the disadvantages that 
post-mortem destruction is not a very safe guide to destruction 
in vivo and that the average quinine-content of a tissue is not 
necessarily representative of all the cells of that tissue. 

It should also be pointed out that so far as extracellular parasites 
are concerned the data showing the quinine-content of various 
tissues, although of much interest in relation to absorption and 
storage of quinine and the physico-chemical conditions inside the 
cells, do not justify any conclusion as to the effective concentration 
of the drug inside the cell—since much of it might be harmless to the 
parasite if precipitated or absorbed by the colloids or dissolved 
in the lipoids of the host cell, or in some non-toxic chemical 
combination. 

As regards the hypothetical places of safety in ‘backwaters’ 
inside the blood-vascular system, it may be pointed out that if 
quinine alone is parasiticidal,. they must satisfy the following 
requirements: — 

(a) Parasites must be able to remain in them throughout such 
periods as the concentration of quinine in the general 
blood stream is actually maintained at a toxic level. 

( 6 ) They must have a blood supply such that even in many 
hours only a small proportion of the total blood passes 
into, them—otherwise quinine would not be present in 
quantity in the general blood-stream so many hours after 
a dose, as in fact it is (cf. (l<Ji8£) page 241). 

(c) They must be surrounded by tissues which remove quinine 
from them at least as rapidly as fresh supplies are 
brought up. 

Of the powers of the various tissues to remove quinine from the 
blood, the data obtained give only imperfect evidence—this power 
must depend not only on the ability of the tissue to destroy or to 
excrete the drug but also on its storage capacity, and in the case of 
those tissues which destroy or excrete quinine this can only be dimly 
guessed at. 

It is, however, clear that the tissues mainly responsible for the 
destruction of quinine are the liver, intestinal wall, muscles and 
kidneys, and that the blood and spleen have little or no such power. 
It is clear, also, that each of the tissues examined, except the lymph 



*74 


glands and red blood corpuscles, store much more quinine weight 
for weight than does the blood or the blood plasma, and that this 
storage capacity is especially high in the suprarenals, spleen and 
kidney. 

The considerable variations in relative storage capacity of the 
different tissues in different experiments (cf. suprarenals with 
intravenous and intraperitoneal injections) may possibly depend on 
variations in the physiological condition of the tissues and in their 
blood supply. 

Failing exhaustive anatomical knowledge of the ‘backwaters’ 
which may exist in various regions of the blood vascular system, and 
therefore considering only the bone-marrow and the spleen in.which 
their existence is beyond question, it is clear that since neither of 
these tissues is capable of destroying quinine post-mortem, such 
quinine-removing power as they possess probably depends solely on 
their storage power, and of this nothing is known except that it is 
greater than in many other tissues and might conceivably be 
adequate for maintaining quinine-free ‘backwaters’ (especially, 
perhaps, in the bone-marrow) provided the access of fresh blood is 
sufficiently slow. 

The lymph-glands are remarkable as being the only tissue in 
which the quinine concentration is consistently less than that in the 
blood. Whether this is due to poor blood supply or to relative 
impermeability to quinine is uncertain. That it is probably not due 
to quinine destruction is indicated by the absence of such in the one 
post-mortem experiment made. 

If the parasiticidal effect of quinine in malaria be not due to 
quinine itself but to one or other of its metabolites, other possibilities 
arise for the survival of the parasite, e.g. they might be safe in 
any backwater if the toxic concentration of the effective metabolite 
is maintained only in the blood of the quinine-destroying 
tissues. Until the existence of such metabolite has been actually 
demonstrated, further discussion would however scarcely be profit¬ 
able. 

That the active metabolite, if it exists, is neither quitenine nor 
any of its metabolite products, is strongly indicated by the observation 
that this substance had no therapeutic value in malaria and was not 
excreted as such in the urine, and by Kemer’s (2) observation that it 



175 

has little or no toxicity for animals although readily absorbed by 
them. It would appear that with the oxidation in the liver of the 
‘vinyl’ group of quinine (R.CH : CH a ) into the carboxyl group of 
quitenine (R.COOH) all therapeutic activity is lost. 

Whether the whole of the quinine which is metabolised in the 
body is converted into quitenine can only be proved by quantitative 
work. If it is, the only metabolite of quinine which could 
conceivably be therapeutically active would be the intermediate 
aldehyde R.CHO. The unknown substance found in ‘hepatised 
quinine’ (Experiment 3, p. 168) is of special interest in this 
connection, and deserves further investigation. 


SUMMARY 

1. Evidence is given bearing on the possibility that there exist 
in the blood-vascular system, regions kept almost free from quinine 
throughout a period of quinine medication. 

2. The quinine content of tissues has been more extensively 
studied. Accumulation at much higher concentration in most tissues 
than in the blood is confirmed. 

3. The suprarenal body is pre-eminent in this respect, although 
not so markedly with intramuscular as with intraperitoneal 
injections. Fairly large accumulations may occur also in the spleen 
and kidney: the lymph glands contain much less quinine than the 
surrounding blood. 

4. The liver, kidney, muscle, intestinal wall, and probably 
pancreas, have considerable power to destroy quinine post-mortem, 
and, therefore, presumably during life. The blood, spleen, supra¬ 
renal bodies, bone-marrow, lymphatic, salivary, and thyroid glands 
have little or no such power. 

5. The quinine-destroying agent extracted from the liver is 
thermolabile, inactivated at ioo° C. and acts best in neutral media. 
Its action is rapid at first, but soon falls off. It does not act at all 
in the absence of oxygen, and is hindered by hydrogen peroxide. 
It can be crudely ' purified ’ by fractional precipitation with alcohol. 

6. Quitenine is formed by the action of liver pulp on quinine. 

7. Quitenine given by the mouth, in ample doses to a malarial 



176 


patient, was therapeutically inert. As none appeared in the urine, 
this is probably true also of its metabolites. 

8. In the faeces of a case of blackwater fever, a brown pigment 
is described which, although not itself cholecyanin, readily yields 
this body. 

9. New tests for quitenine and quinotoxin are described. 

10. By an improved procedure, the thalleioquin test is rendered 
capable of detecting easily 0*004 mgm. of quinine. 

In conclusion, I wish to record my thanks to Professor W. 
Ramsden for the valuable help and advice he unstintingly gave me 
during this work, and for his collaboration in discussing the results 
of the investigation. 

I am indebted also to Professors J. W. W. Stephens and 
W. Yorke for help with clinical experiments and animal injections. 


REFERENCES 


Giemsa, G., and Schaumann, H. (1907). Arcbiv. f. Scbiffs. u. Tropen. Hygiene , Beiheft. 3. 

Keener, G. (1870). Beitrage zur Kenntruss der Chininresorptin. Arcbiv. f. d. gesammte 
Physiologic. 

Merkel, A. (1902). Stoffwechselproducte des Chinins. Arcbiv. f. Exp. Path. u. Pbarm Vol. 
XLVII. 

Nierenstein, A. M. (1919). * Quitenine,* a disintegration product of quinine found in urine. 

Jovrn. Roy. Army Med. Corps, VoL XXXII, pp. 218-219. 

Ramsden, W., and Lipkin I. J. (1918). Detection and estimation of quinine in blood and urine. 
Ann. Trap. Med. & Parasitol ., Vol. XI, pp. 434-464. Brit. Med. Jottrr. ., May, 1918. 

Ramsden, W., Lipkin, I. J., and Wfitley E. (1918). On quinine in animal tissues and liquids 
with methods for its estimation. Ann. Trop. Med. &? Parasitol Vol. XII, pp. 223*258. 

Skraup (1880). Monatsbefte fur Cbemie , Vol. X, p. 39. Bericbte d. deut. Cbem. Cesell , Vol. 
XII, 1104. 



'77 


CASES OF ACUTE AMOEBIC DYSENTERY 
IN ASYLUM PATIENTS NEVER OUT OF 

ENGLAND 

BY 

A. MALINS SMITH, M.A., Cantab. 

(Received for publication 6 June , 1919) 

In a previous paper issued from this Laboratory, Matthews and 
Smith (1919) recorded the results of the protozoological examina¬ 
tions of the stools of two hundred and seven patients at 
Whittingham Asylum. These stools were examined as fresh as 
possible, yet, as they were not examined at the asylum itself, several 
hours (in actual practice from about eight to about thirty) necessarily 
elapsed before the examinations could be made. Through the kind 
introduction of Professor Warrington Yorke, I was recently enabled 
to carry on examinations at an asylum itself, namely the Lancashire 
County Asylum at Rainhill. In these circumstances the stools could 
be examined very soon after being passed. Care was taken that 
the dysenteric stools in particular should be examined as fresh as 
possible. Some were examined within half an hour of being passed, 
and five hours was the longest time which elapsed before a dysenteric 
stool was examined. My visit coincided with a considerable 
outbreak of the so-called ‘asylum* dysentery, and I had the 
opportunity of examining, during my stay of some two months, the 
stools of sixty patients suffering from acute dysentery. In three of 
these patients I found free amoebae containing ingested blood 
corpuscles, i.e. Entamoeba histolytica . 

It has previously been considered that asylum dysentery was of 
bacillary origin. Dr. Gettings (1915), Pathologist at Wakefield 
Asylum, isolated, at the height of the 1913 epidemic, Flexner’s 
bacillus from 50 per cent, of the cases he examined. This 
leaves half the cases unaccounted for, and by no means precludes 
the possibility of some of the cases being amoebic. In the 
discussion on Dr. Gettings* paper, it was stated that ipecacuanha 



i 7 8 


was used very largely by the visiting physicians at Wakefield 
Asylum, nearly a century ago. From this fact, and from the fact 
that relapses occur in asylum cases, it was suggested that some of 
the Wakefield cases might possibly have been amoebic and the 
possibility still remains, as Dr. Gettings confessed that he had not 
examined for the amoeba. 

RESULTS OF EXAMINATIONS OF THE RAINHILL 
PATIENTS 

i. The acute cases 

.1 have included among these every case in which blood and pus, 
or pus alone; or blood and mucus were present in the stool. I have 
not included those cases with blood alone, which might have had 
other origin, e.g., haemorrhoids, nor with mucus alone, as mucus is 
commonly present in colitis and post-dysenteric conditions. 
I examined the stools of sixty patients showing acute dysentery, as 
thus defined, and of these 3 (5 per cent.) showed the free Entamoeba 
histolytica containing red blood corpuscles. Two of these patients 
(males) had never been out of England, and one (a female) had 
lived in Rouen during girlhood, but had been nowhere else outside 
this country. The occurrence of these cases is of considerable 
interest, and I, therefore, append some details of each case. 

J. Ho., aged 39, male, admitted February 18th, 1918, from 
Wakefield Asylum, admitted there July 21st, 1917, from Pontefract, 
his native place; was never abroad, was in poor health on 
admission with cardiac and kidney disease. His stool was first 
examined on April 9th, 1919, when it was normal and negative. 
On April 13th he developed symptoms of dysentery, and an examina¬ 
tion of his stool on April 14th revealed the presence of blood, mucus 
and pus and Trichomonas flagellates. On April 17th his stool was 
again examined, and large numbers of motile amoebae containing 
red blood corpuscles were present in the bloody mucus. There 
were present also heavy infections of Trichomonas and Chilomastix . 
He was given half a grain of emetine hydrochloride subcutaneously 
twice daily on the 20th and 21st April. On the 22nd his stool was 
negative. The treatment was accidentally discontinued from April 
22nd to 30th, and his stool was positive on April 23rd. A twelve 
days’ course of emetine, as before, lasting from May 1st to 12th, 



*79 

resulted in gradual improvement. The highest temperature during 
the illness was ioo'4° F. 

J. Ha., aged 32, male, admitted December 20th, 1912, born in 
Scotland, never abroad, lived lately in Warrington, no illness at 
Rainhill until present attack, but had some symptoms of kidney 
disease. On April 17th, 1919, the first examination of his stool was 
made. He was found to be passing cysts of Entamoeba histolytica. 
The stool was normal. On April 25th he developed symptoms of 
dysentery and had mucus, pus and blood in his stools, which were, 
however, negative for protozoa. On April 28th amoebae were 
present in the stools. Owing to the absence of erythrocytes these 
could not be diagnosed. On April 29th, however, amoebae with 
ingested red blood corpuscles were seen. A course of 12 grains of 
emetine hydrochloride, subcutaneously, 1 grain per day, was given 
from May 1st to 12th. This caused immediate marked improve¬ 
ment in the number and consistency of the stools, and the patient 
soon completely recovered. Highest temperature 99*4° F. 

L. P., aged 37, female, admitted 21st June, 190P, bom near 
Liverpool, was always delicate, but no definite illness, lived in Rouen 
for some time when a little girl, no known illness there. No other 
journey abroad. Her stool was examined once only on April 29th, 
1919, when blood, mucus and pus were present and also amoebae of 
E. histolytica with ingested red blood corpuscles. A course of 
emetine hydrochloride was given from May 2nd to 13th. The 
number of stools passed in the first week of her illness was thirty-five 
(April 29th to May 5th). From May 5th to 12th, while the emetine 
was being given, the number was reduced to sixteen, and in the 
following six days it was fifteen, so that marked improvement 
resulted. Highest temperature 10I‘6°F. 

Thus the fact of acute amoebic dysentery occurring in an asylum 
is definitely established. No exactitude can be claimed for the 
figure—5 per cent.—showing the proportion of amoebic to all acute 
cases. Possibly it may be higher than this in other asylums, and it 
is, even at Rainhill, almost certainly a minimum. As the examina¬ 
tions of the case J. Ho. showed (cf. Wenyon and O’Connor (1917), 
pp. 65 and 66), even acute cases do not pass amoebae in every stool, 
and in the great majority of the cases only one stool was examined. 
There seems to be no doubt, however, that the majority of the cases 



i8o 


were bacillary in origin. It is commonly stated, see Wenyon and 
O’Connor (1917) and Willmore and Shearman (1918), that a great 
preponderance of pus in the Stools, as seen under the microscope, 
is a characteristic of bacillary dysentery. I noted those cases in 
which pus was the most abundant constituent on microscopic 
examination, and these numbered thirty-six out of fifty-nine, 
i.e., 60 per cent. In these thirty-six no amoebae were found, the 
three amoebic cases being found among the twenty-three cases in 
whose stools pus was a minor constituerit or altogether absent. The 
preponderance of stools in which pus was the most important 
constituent and the finding of amoebae in only a small proportion of 
the cases renders it probable that the outbreak was mainly bacillary. 
It will be noted later (see Table IV) that the percentage of carriers 
of E. histolytica in the asylum was smaller than that found in 
Whittingham Asylum. It is probable, if an outbreak of dysentery 
should occur in an asylum with a higher percentage of E. histolytica 
carriers, that the percentage of acute amoebic cases would be higher 
than at Rainhill. One aspect of the case J. Ha. may, perhaps, 
be emphasised in this connection. He was a carrier with normal 
stool on his first examination, and according to present views (see 
Dobell and Stevenson (1918), but see also Matthews (1919)) had 
probably been a carrier for a considerable period. Yet his 
dysenteric symptoms did not appear until the height of an outbreak 
of dysentery which was undoubtedly mainly bacillary in origin. No 
bacteriological examinations of his stools were made, and it is possible 
that the primary cause of his attack was a bacillary one. A case of 
dysentery whose exciting cause was Flexner’s bacillus, for example, 
would probably pass E. histolytica amoebae, if the patient was 
already a carrier of E. histolytica. In such a case E. histolytica 
might only cause a secondary aggravation of the condition. In the 
absence of bacteriological examinations no definite statement on this 
point can be made. 

As bearing on the question of the place of origin of the acute 
dysentery cases, it may be noted that some of the cases were 
patients who had recently been admitted to the asylum. Of the fifty- 
nine* acute cases, fourteen were patients admitted within the three 

• One of the 60 acute case# previously mentioned was found at post-mortem to be a case 
of carcinoma of the rectum. 



i8i 

months previous to the onset of their attack. Seven had dysenteric 
stools within one month of their arrival in the asylum, and of these 
four had never been abroad, while three had been in the tropics. 
Five showed symptoms of dysentery within a fortnight after 
admission (the exact intervals being fourteen, twelve, six, five and 
one day respectively), and of these three had never been abroad, 
while two had visited the tropics. It cannot, of course, be in any 
case proved that the disease originated outside the asylum, but it 
seems probable from the figures that the asylum receives from the 
outside population, both from those who have been in the tropics 
and those who have never been abroad, occasional cases of dysentery. 

2. The general population of the asylum 

The total number of asylum patients examined, inclusive of the 
acute cases, was five hundred and four. The majority of these 
patients were without any record of physical ailments. The 
commonest ailment was dysentery, and sixty-nine of the patients 
were suffering from, or had a record of, this disease. Twenty-five 
had renal disease. Nine of these twenty-five had also acute 
dysentery. In two of these nine cases the dysentery was amoebic. 
Other illnesses, apart from mental trouble, were comparatively rare. 
The average age of the patients examined was 42 years. 

Table I gives the result of the examination of these five hundred 
and four patients for intestinal protozoa. The figures are based on 
one examination per case. 

The'stools of a few of the patients were examined more than 
once, and, as usual, new infections were found at later examinations 
than the first. Sixy-two patients received two examinations each, 
eleven three each, three four each, and two five each. The examina¬ 
tions subsequent to the first resulted in finding new infections as 
follows:— Entamoeba histolytica , four; E. coli, seven; E. nana, 
three; Giardia intestinalis, four; Chilomastix mesnili, six; Tricho¬ 
monas intestinalis, two; and ‘ I ’ bodies, one. The results of these 
later examinations have not been included in any of the tables given 
in this paper. 

In addition to the infections recorded in Table I, five cases with 
‘ I ’ bodies (ro per cent.) were discovered, and also Oxyuris eggs in 
one case, Taenia sp. eggs in two cases, and Trichuris eggs in one 



case. As was to be expected, there is no significant difference 
between the infections of the men and those of the women. Giardia 
intestinalis is comparatively rarer in the latter, but a larger number 
of cases would be necessary before any importance could be attached 
to this difference. 

An attempt was made to ascertain if there was any significant 
increase in the number of infections as the length of stay of the 
patients increased. The difficulty at once presented itself that sub¬ 
divisions of such a—for statistical purposes—comparatively small 
number as five hundred are too small to give significant results. It 
may be worth while to note, however, that there were sixty-three 
patients examined who entered the asylum after January 1st, 1919 


Table I. 



Number 

examined 

Entamoeba 
histolytica 
infections 
per cent. 

E. coli. 
per cent. 

E. nana. 
per cent. 

Giardia 
intestinalis 
per cent. 

Cbilomastix 
mesnili 
per cent. 

Tricho¬ 
monas 
intestinalis 
i per cent. 

Men. 

! 

285 | 

3*5 

1 

i 

! 20* I 

B 

67 

6*2 

07 

Women 

219 

1 5 -o 

| 22*8 


*7 

7*3 

n 

Total 

504 

4*2 

21*4 

■ 

5 *° 

n 



(the examinations were carried on from April 7th to May 15th, 1919), 
and among these sixty-three, there occurred one infection with 
Entamoeba histolytica, ten with E. coli, three with E. nana, and one 
with Giardia intestinalis. Thus, as was to be expected, infections 
with the various protozoa are being received from outside. The only 
other figures on this point which I have thought worth recording 
are found in Table II, where the five hundred and four patients 
examined have been divided into two almost equal groups according 
as they have been more than or less than four complete years resident 
in the asylum. 

The significance of these figures is at present very difficult to 
estimate, and I shall not attempt to draw conclusions from them. 

Every stool examined was seen in bulk, and the presence or 
absence of mucus and blood noted. The stool was further classified 
as loose, semi-formed or formed. The formed stools numbered 












Tablk II. 



Number 

examined 

Entamoeba 
histolytica 
per cent. 

E '. coli. 
per cent. 

E. tuna. 
per cent. 

Ciardia 
intestinalis 
per cent. 

Cbilomastix 
mesnili 
per cent. 

Tricho¬ 
monas 
intestinalis 
per cent. 

Resident less 
than four 
years 

249 

2*0 

21*6 

36 

■ 

5*2 

0*4 

Resident 
more than 
four years 

*55 

64 

21*2 

1 ‘ 4 

4 ‘o 

8*2 

1*2 


exactly half of the total number examined. With the exception of 
one case of flagellate trichomonas in a formed stool, the free forms of 
the flagellates and of the entamoebae were found in the loose or semi- 
formed stools only. 

Table III shows the percentage of the intestinal protozoa found 
in the formed stools on the one hand and the loose and the semi- 
formed stools on the other. The only figure worthy of note is that 
for G. intestinalis. The larger number of infections of this flagellate 
found in the loose stools agrees with the results of Wenyon and 
O’Connor (1917) on this point. 


Table III 



1 

1 

Number 

examined 

Entamoeba 
histolytica 
per cent. 

E. coli . 
per cent. 

E. nana. 
per cent. 

Ciardia 
intestinalis 
per cent. 

Cbilo¬ 
mastix 
mesnili 
per cent. 

Tricho¬ 
monas 
intestinalis 
per cent. 

Formed stools 

252 

48 

240 

3*6 

n 

7*2 

! °*4 

All others ...1 252 

i 

l 

3*6 

19*2 

2*4 

6*8 

64 

1*2 


A note may be made of the fact that in one of the men’s wards, 
from which the stools of sixty-nine patients were examined, the 
percentage of G. intestinalis was found to be 14^ and that of 
C. mesnili ii|. These figures, particularly the former, are 
significantly larger than those for the whole number examined. The 
variation may be due to chance, as the number examined was not 
large, but on the other hand it may be correlated with the fact that 
this ward contained, in the words of Dr. Cowen, the Medical 












184 

Superintendent, * patients who are a very degenerate lot and are in 
many cases of faulty habits.’ 

Finally Table IV gives the results for Rainhill, compared with 
other sections of the population already examined and reported on. 
As only thirty-four of the five hundred and four patients at Rainhill 
had, so far as could be traced, ever been abroad, the figures are to 
be looked upon as representing the infections of a section of the home 
population, and are to be compared with the groups of that popula¬ 
tion rather than with the results for returned soldiers. 


Table IV. 




Home Population 


Returned Soldiers 


Rainhill 

Asylum 

Whitting- 

ham 

Asylum 

Adult 

Civilians, 

Hospital 

population 

Children, 

Hospital 

population 

Army 
Recruits j 

1 

Dysenteric 

Convales¬ 

cents 

t 

Ndn- 

dysenteric 

Convales¬ 

cents 

Number 

examined 

5°4 

207 

450 

5+8 

1,098 

4,068 

45 ° 

Entamoeba 

histolytica 

4*2 

97 

*’5 

r8 

5-6 ! 

r° 

64 

E . coli. 

21*4 

45*9 

67 

1 ri 

18*2 

15-2 

14*2 

E. nana. 

3 ’° 

12*1 

2*4 

27 

5*5 

... 


Ciardia 

intestinalis 

5 '° 

3‘4 

6*o 

14-1 

7-0 1 

9*9 

6’o 

Cbilomastix 
mesnili ... 

67 

23*2 

r 5 

r8 

i 

0*2 

i 

3*6 

(3 exams, 
per case) 

2*0 

(2 exams, 
per case) 


It may be noted that the various protozoal infections, with the 
exception of G. intestinalis, are much less common in the patients at 
Rainhill Asylum than we found them to be among those at 
Whittingham. It may be that wide differences would be found in 
the various asylums in this respect, differences depending on factors 
at present unknown. The comparative isolation of an asylum 
population would tend to prevent these differences, once established, 
from being obliterated. Though the number of G. intestinalis 
infections is higher than at Whittingham, yet the difference is not 
so marked as to call for any revision of the theory put forward 
in previous papers by Matthews and myself, namely that this 



flagellate tends to disappear from older people. In fact, the smaller 
percentage of G. intestinalis among those who have been longer in 
the asylum (see Table II) would form some slight further evidence 
in favour of our view. 

I wish to thank very heartily Dr. Cowen, Medical Superintendent 
at Rainhill and Dr. Watson, Pathologist, for their great help and 
kindness during the course of my investigation. But for their 
cordial co-operation the work would have been impossible. 


SUMMARY 

Five hundred and four asylum patients have been examined for 
intestinal protozoa. Fifty-nine of these had acute dysentery and in 
three of the cases vegetative E. histolytica were found in the stools. 

All the usual protozoa were found in the stools, but the infections 
were distinctly less numerous than in the patients of the asylum 
previously examined. 


REFERENCES 


Gettings (1915). Bacillary Dysentery. Irani. Trop. Med. and Hyg ., Vol. VIII., p. in. 

Winyon and O’Connor (1917). Human Intestinal Protozoa in the Near East. Wellcome 
Bureau of Scientific Research , London. 

Willmore and Shearman (1918). On the differential diagnosis of the dysenteries. Lancet 
CXCV., 2, p. 200. 

Dobell and Stevenson (1918). A note on the duration of infections with Entamoeba histolytica. 
Trans. Trop. Med. and Fyg., Vol. XI., pp. 168-175. 

Matthews (1919). The course and duration of an infection of Entamoeba coli. Ann. Trop. Med. 
and Parasitol ., Vol. XIII., p. 17. 

Matthews and Smith (1919). The spread and incidence of intestinal protozoal infections 
in the population of Great Britain. IV. Asylum patients. V. University and school 
cadets. Ann. Trop. Med. & Parasitol., Vol. XIII, p. 91. 




■87 


THE EXPERIMENTAL INFECTION, IN 
ENGLAND, OF ANOPHELES PLUM- 
BEUS, Hal., WITH PLASMODIUM 
EIFAX. (SPOROZOITES IN SALIVARY 

GLANDS.) 

(Preliminary Note) 

BY 

B. BLACKLOCK 

AND 

H. F. CARTER 

(.Received for publication 24 July, 1919) 

Specimens bred at the Liverpool School of Tropical Medicine 
from the pupae, collected in England, of A. plumbeus, Hal., were 
fed on a volunteer patient, suffering from simple tertian malaria, 
acquired in Salonica. On 9.7.19 the patient had a rigor at 9 p.m. 
and the blood was positive; on 10.7.19 at 6 p.m. the blood of the 
patient showed a moderate infection of trophozoites and gametes. 
The Anopheles were now fed on him, and six of them which had 
fully engorged were separated and placed in glass globes in an 
incubator, kept at a temperature of about 28° C. The mosquitoes 
received no meal of blood either before or after the infective feed, 
but were given raisins. The first mosquito died on 13.7.19, the 
second on 14.7.19, the third on 15.7.19; no evidence of infection was 
found in any of them, but the presence of blood in the gut rendered 
the examinations unsatisfactory. The fourth mosquito, however, 
which died on 18.7.19 was found to be infected, twelve oocysts being 
found in the gut, some of them in an advanced state of development. 
The fifth mosquito was killed on 21.7.19 at 10 a.m., and infections 



of both the gut and salivary glands were found. Both glands were 
infected with sporozoites in large numbers, the distal extremities of 
the lobes being chiefly involved. 

We have also obtained oocysts (thirty-six, varying in size from 
18 At to 30/1) in the gut of a female killed on the thirteenth day after 
the infective feed, and kept at laboratory temperature. 



189 


NOTE ON A CASE OF MULTIPLE 
INFECTION BY DRACUNCULUS 
MEDINENSIS 

BY 

B. BLACKLOCK 

AND 

Capt. W. R. O'FARRELL, R.A.M.C. 

Plate VII 

(Received for publication 22 July , 1919) 

The following case of guineaworm infection presents some 
features of interest, namely the number of worms with which the 
patient was infected, and the somewhat severe symptoms caused by 
their presence and condition. 

The patient, an Indian, aged 15, from Nizampur village in the 
district of Alibag, was admitted to the Tropical Ward of the Royal 
Infirmary, Liverpool, on June 3rd, 1919. He stated that he had 
only been away from India about two months, that neither he nor 
the members of his family had suffered from this disease, but that 
it was common in his village. 

Local condition . Both feet and the right hand were affected, 
the lesions being distributed as follows: — 

Right foot: (1) Immediately above the internal malleolus; 
(2) below and behind the internal malleolus; (3) the outer side of the 
first phalanx of the second toe. 

Left foot: (1) On the dorsum of the fourth metatarsal; (2) below 
and behind the external malleolus; (3) below and two inches in front 
of external malleolus; (4) on the inner aspect of the dorsum of the 
first metatarsal. 

Right hand: (1) On the radial side of the base of first meta¬ 
carpal; (2) on the dorsum of the first phalanx of the middle finger. 



190 


In all these sites the lesions were of much the same character. 
The superficial layers of the skin had been removed over wide areas, 
especially of the feet, probably by previous applications. The 
lesions caused by the worms consisted of a more or less circular 
area—measuring from 2 to 3 centimetres in diameter—of raised 
tissue with a central orifice. In the majority of the lesions the tissue 
round the opening was scaly and white, but in two there was a 
prominent area of red granulation tissue extending 1 centimetre 
around the orifice. From several of the sites portions of worm of 
varying length were protruding, and from all the sites, by gentle 
pressure, a small amount of pus could be obtained. Both feet and 
the right hand was swollen, oedcmatous, and tender. 

The appearance presented by the patient’s feet and hand are 
shown in Plate VII, figs. 1-4. 

Among the portions of worm protruding from the openings, 
no head was discovered. They had been removed previous to 
admission. Examination of the discharge from the lesions showed 
that although larvae were present, they were motionless. Larvae 
removed from the body of the adult females also proved to be dead. 

General condition. The temperature was 98 4° F., pulse 94 0 . 

Blood. No parasites were observed either in fresh or stained 
preparations. Eosinophilia was noted to the extent of 14 per cent. 
(500 leucocytes counted). 

Urine. Clear, sp. g. 1009, acid, no alb., blood, or sugar. No 
deposit macroscopic or microscopic. 

Faeces. There was a small quantity of apparently pure blood 
passed with the first motion, but this was possibly present owing to 
the scybalous nature of the motion. It was not observed on subse¬ 
quent examinations. Ova of Ascaris lumbricoides and Trie kurus 
trichiura were found, and later ancylostome ova also. 

Treatment. In view of the condition of the lesions, neither 
gradual extraction of the worms nor the injection of them or the. 
tissues with a view to preventing the spread of sepsis appeared to 
hold out much chance of success. It was considered possible that 
the intravenous injection of antimony might be of some value in this 
direction. On the 6th and 8th June the patient received an 
injection of I grain of tartar emetic. On the yth June the patient 
had some epistaxis and coughed up some blood-stained sputum, but 



there is no evidence that this was due to the injection, and no such 
symptoms followed the second injection. On the 7th of June the 
portion of worm presenting on the right wrist disappeared from 
view; on the 8th, three inches of worm were discharged from the 
left foot; on the 9th, fresh lesions were observed, one on the right 
wrist, from which a portion of worm emerged, and another on the 
left foot; also five inches of worm were discharged from the right 
foot; all the openings were discharging freely pus tinged with blood. 

At this stage the rapid progress of the septic condition accom¬ 
panied by enlarged and painful glands in the groin necessitated 
surgical interference. It was, therefore, not possible to come to any 
conclusion as to the effect of treatment by intravenous injection of 
antimony in this case of guineaworm disease, as the case did not 
afford the drug a fair trial. During this period the patient’s 
temperature rose nightly to over I02°F. (see chart), his pulse and 



respiration also being rapid. X-ray examination of the limbs was 
negative. On June 12th, Mr. Thelwall Thomas opened up the worm 
tracks in the feet, and removed portions of several worms, ligatures 
being applied to the portion remaining in the tissues. No tail 
portions were obtained, but the next day at dressing the terminal 
portions of two worms were removed easily, one from the left foot, 
eight inches in length, and one from the right hand, each bearing 
the typical curved tail. Further terminal portions of tails were 
obtained at subsequent dates. The following table shows the period 
during which the portions of worms were obtained: — 





192 


Table 

Giving total lengths of portions of worms obtained from each limb, with the number of 

tails found. 


Date 

Limb 

Total length of 
portion! in inches 

Tails 

(1) From 8.6.19 t0 19-6.19. 

Left foot 

49 

3 

(2) From 13.6.19 to 15.6.19 

Right hand 

16 

2 

(3) From 5.6.19 to 7.19 . 

Right foot 

57 

3 

Total 


122 

8 


On 25.6.19 the patient had a rigor, temperature io5°F., but no 
malarial parasites were found on several examinations. Quinine 
was given and appeared to exercise some effect on the temperature. 
The right foot was tender and swollen. On 3.7.19, under a general 
anaesthetic, the right foot was further incised and a portion of worm 
extracted, bearing a tail. 

The larvae. As stated above, these were all motionless, and 
they did not recover in water. The average length of fifty, drawn 
and measured, was 616/u, maximum 737/», minimum 490/1. In 
transverse section of the female worm the larvae were, in the majority 
of cases, cut transversely, some more obliquely. The larvae in 
section stained readily, whereas, when free, they proved difficult to 
stain. 












EXPLANATION OF PLATE 


Showing the appearances presented by the feet. 
Showing the appearance presented by the right hand. 



4finals Trop. Med . & ParasitolVol. XI!I 


PLATE VII 










x 95 


COINCIDENT MALARIA AND ENTERIC 

FEVER 

BY 

HENRY HAROLD SCOTT, M.D., M.R.C.P., London; 
F.R.S.E., D.P.H. 

government bacteriologist, Jamaica, b.w.i. 


(Received for publication 17 June , 1919) 

1 Twenty years ago, when it was alleged before the Indian 
Plague Commission that anti-plague inoculation had cured eczema, 
gonorrhoea, and other miscellaneous infections, I thought the matter 
undeserving of examination. I took the same view when it was 
reported in connection with anti-typhoid inoculation that it rendered 
the patients much less susceptible to malaria.* These words were 
spoken by Sir Almroth Wright (1919) in the course of a paper read 
at a meeting of the Royal Society of Medicine and reproduced also 
in the Lancet . 

During the last few months, in a report to the Tropical Diseases 
Research Committee, I mentioned certain peculiarities which were 
noticeable in cases of coincident enteric fever and malaria, and as 
having some bearing on the second of the above two passages the 
following account may be of interest. 

Every worker in the tropics has met with cases of enteric fever 
in which malaria supervenes, but in the majority, if not in all, of 
these about to be described both conditions were present together. 
Lyon has published some cases, but his paper is not available, and 
I cannot tell, therefore, whether his experiences coincide with my 
own. Far be it from me to attempt to revive the old term of 
* Typhomalaria,’ for unless this is distinctly understood to imply 
merely malaria occurring in a typhoid patient, the term is liable to 
4 mislead in giving the idea of a single and distinct disease. 
When, however, we have a dual infection from the beginning, certain 
peculiarities manifest themselves which are worthy of special remark. 



196 


It has been stated by Sir Patrick Manson that in malarious 
subjects the oncoming of typhoid is often preceded by three or four 
paroxysms exactly like those of ordinary ague. In such cases 
quinine is usually given early, and its failure to check the disease 
should warn us to be guarded in giving a prognosis. Vice-versd, 
well marked malaria-like fluctuations of the temperature and the 
appearance of parasites in the blood in the course of a continued 
fever do not exclude typhoid. These cases are ‘ typhomalarial,’ that 
is typhoid fever with a malarial complication. 

One must remember in discussing this question that any disease 
in a malarial subject is prone to take on an intermittent or periodic 
character; thus one meets with cases of true lobar pneumonia, for 
example, with a distinctly periodic remission until the crisis takes 
place; this must not be confused with the ‘ pneumonic form of 
malaria,’ such as occurred in some of the Salonika troops. 

No attempt will be made to discuss the question of the simulation 
of malaria by typhoid nor that of typhoid by malaria, but that of the 
definite co-existence of the two conditions. There are, of course, 
three periods at which this combination may take place. Firstly, 
at the onset; in such cases the usually described ‘Typhoid chart’ is 
departed from. The rise is not of the staircase type at all, but with 
rigors, high fever, sweating, partial remissions, strongly suggestive 
of and usually diagnosed as malaria, especially as similar attacks 
may be repeated for two or three days with, it is true, smaller 
remissions as a rule. Quinine is given, the rigors and marked 
oscillations are checked, but the temperature remains persistently 
high and the typical typhoid condition develops. 

Secondly, the attack begins in the customary insidious manner of 
most cases of enteric fever, then, about the second or third week, 
rigors and oscillations occur giving rise to considerable anxiety as 
to urgent complications. Administration of quinine brings about a 
cessation of these oscillations, and the typhoid resumes its course. 

Thirdly, the malaria may occur, as already stated, at the 
termination of the enteric fever attack or during convalescence. In 
some of these cases one sees in a patient suffering from Plasmodium 
vivax infection fluctuations of temperature strongly suggestive of 
tertian ague (though, as stated, rarely falling actually to normal), 
but there are the furred tongue with red tip and edges, the loose 



*97 


pea-soup stools, the listlessness and other clinical appearances of 
enteric fever. Again, with concomitant P. falciparum and enteric 
infection the chart may be one of subtertian malaria with inter¬ 
missions, but the appearance clinically is that of an enteric patient. 

The following are brief descriptions of some cases recently met 
with: — 


1. P.Y., male, aged 28 years. 

Was admitted with a temperature of ioi° F.; previous duration uncertain ; 
was said to have 1 had fits a few days ago,’ before coming to hospital, but nothing 
definite as to the nature or character of these could be ascertained. His chart 
showed that during his stay in hospital the highest temperature was 101*4° F. 
One week after admission it fell to normal for 24 horns, and four days later reached 
normal to stay there. The debility was but slight, and recovery rapid and unevent¬ 
ful. On the day of admission the blood was examined and gave an agglutination 
of B. typhosus in high dilution and also of B . paratyphosus a. in low dilution (1 : 30 
only), probably a group reaction ; P.jalciparum was seen in smears. The malarial 
element is clear; the question is whether there had been an old attack of typhoid 
fever leaving residual agglutinins in the serum. The titre was high, so that if 
there had been a previous attack the patient could hardly have forgotten it, yet 
no history of any prolonged fever could be obtained. 

2. J.W., male, aged 12 years. 

This patient gave a history of ‘ fever and headache * for three weeks prior to 
coming to hospital; on admission the temperature was ioi° F., and the tongue 
was coated. Subtertian parasites were found in the blood and quinine was given. 
In view of the history and of the facts that the temperature in four day s, though 
lower, had not become normal, and that the patient appeared ill, a Widal reaction 
was asked for and a strong agglutination of B. typhosus was obtained. So much 
of the illness as was observable, namely, at the end of the third week and after, 
showed a temperature like that of a mild case of enteric fever at that period. 
The malarial infection did not appear to have any appreciable effect on the 
temperature. He had had no quinine or any other form of medicinal treatment 
before coming to hospital. Ten days afterwards the temperature reached normal 
and stayed there; convalescence was rapid and uninterrupted. 

* 3. B.M., female, aged 21 years. 

This patient was admitted to hospital with a history of ‘ fever and headache ’ 
for the preceding four days; there had been no shivering attacks. Her temperature 
was 102° F. on coming to hospital. Blood examination revealed the presence of 
P.jalciparum ; the serum gave a marked agglutination of B . typhosus in low dilution 
(up to I in 50), but blood culture yielded a growth of B. typhosus . The tongue 
was coated, with red tip and edges, and the general aspect was that of an enteric 
patient. In view of the presence of malarial parasites quinine was given, but 
otherwise the treatment was that of enteric fever. During the succeeding five 
days the temperature once fell to 99° F., but except for this remained between 
102 0 and 104°. On the ninth day of illness it fell rapidly to 98°, and remained 
normal till the day of discharge from hospital three weeks after admission (Chart 1). 



198 



This is interesting because, if, as would appear to be the case, 
there was definite infection with B. typhosus , the attack was 
exceptionally mild. No previous history of any prolonged fever 
could be obtained whereby the Widal result might be regarded as a 
residual agglutination reaction. Moreover, the patient presented a 
typhoid appearance, and the positive result occurring in compara¬ 
tively low dilution (1 : 50 only) would be accounted for by the early 
stage of the disease, the fifth day. Other possible explanations are 
either that the attack was abortive (by no means synonymous with a 
mild attack), or what is less likely, the patient was not actually 
suffering from an attack of enteric fever, but that the agglutination 
was given owing to a previous infection, or lastly, that she was a 
1 healthy carrier * without having undergone a previous attack and 
that the presence of the bacilli was merely a temporary, transient 
condition during which one was fortunate enough to obtain a 
culture. This last is a far-fetched explanation, but in whichever 
way the question is regarded there are difficulties of interpretation. 

4. A.W., male, aged 26 years. 

This man was admitted to hospital with a history of * fever, headache, and 
chills ’ during the preceding eight days. The temperature on admission was 
103° F., the tongue furred in the centre, but with red tip and edges. In view 
of the history 9mears of blood were taken in the admission room and examination 
revealed P. jalciparum and pigmented mononuclears. Quinine was given by 
mouth and later, as the temperature remained elevated, by injection in large 
doses. This, however, was not effectual and the Medical Officer, suspicious of 






i 9 9 


‘ something besides malaria being at the root of the trouble,’ sent up the blood 
for agglutination tests. B. typhosus was agglutinated in high dilution. The 
patient had by this time been ill for three weeks, but during the following seven 
days the fever abated, the temperature becoming normal on the twenty-ninth 
day of disease and remaining there (Chart 2). 



Two points to be noticed in this case are, firstly, the rapid fall 
of the temperature to normal to remain there instead of showing the 
usual terminal fluctuations of a case of enteric fever; secondly, the 
combination of malaria with enteric did not prove any more 
severe than the latter alone. This fact has been noticed in several 
instances. 

5. R.D., female, aged 17 years. 

Prior to admission to hospital this patient had been ill for eight days with 
‘ fever, pains in the head and abdomen.’ The temperature in the taking-in room was 
found to be 105° F. ; the pulse was 115 ; the spleen was enlarged, and there was 
abdominal tenderness, especially in the right iliac region. The blood was examined 
and P. falciparum found in considerable numbers, while the Widal reaction gave 
a marked agglutination of B. typhosus ; the degree of this was greater in the higher 
dilutions than the lower—a reversed reaction (pro-agglutinoid). Four days later 
there is a note to the effect that the patient was delirious at night, but this subsided, 
and a week afterwards it is stated that 4 she is sleeping well.’ Improvement set 
in with a sudden fall of temperature from 102° to 98° and this was maintained 
for eight days, when, without any return of the fever, the patient became dull 
and apathetic, took nourishment badly, and grew progressively weaker. Malarial 
parasites were not again found in the blood, and there was no further rise of 
temperature till 24 hours before death, which took place on the fifty-sixth day 
of disease. Her condition during the last three weeks of life was that of a post' 
typhoid psychosis, a sort of lethargy (Chart 3). 




1 









200 



Chart 3 


The same point is to be noticed here as was mentioned in the 
previous case. The temperature, io5 °F. on admission, varied 
between 102° and 103° for four days. During the succeeding eight 
days it never exceeded 102°, and at the end of the third week of 
illness dropped rapidly from 102° to 98° and thereafter, except when 
99 0 was registered on three occasions, it remained subnormal till the 
last twenty-four hours of life, during which it rose to ioi°. The 
febrile course is thus seen to have been a mild one, and the presence 
of P. falciparum did not appear to render it any more severe; the 
fever was of an unusual type for the third week of typhoid fever, but 
examination of the blood again during that period did not result in 
the finding of any malaria parasites to account for the irregularity 
or the sudden drop. 

6. Z.P., male, aged 53 years. 

Prior to coming to hospital this patient had for eight days, so he stated, been 
suffering from ‘ fever, pains all over, and cough.’ The temperature on admission 
was ior6° F.; the stools were loose and offensive. Blood examination on day 
of admission : P. falciparum present; the Widal reaction was indefinite : there 
was considerable loss of motility and attempts at agglutination, that is, there were 
small clumps of bacilli, but the majority were isolated though motionless, and 
this was noticed only in the lower dilutions. At the end of the third week of 
illness the temperature became normal. On the fourteenth day of disease the 
patient’s condition was poor, he was very weak, and the pulse soft and feeble, 
but after the fall of the temperature he rapidly improved and convalescence was 
uninterrupted. The blood was again tested for agglutination, and gave a well 
marked positive result with B. paratyphosus A ; negative w'ith B. typhosus and 
B . paratyphosus b. No malaria parasites found in smears. With the exception of a 


X 





















201 


single occasion on which 99 0 was recorded, it will be noticed again in this case that 
after once falling to normal the temperature did not again rise and never showed 
any of the usual terminal oscillations of most enteric patients (Chart 4). 



Chart 4 

7. R.P., male, aged 7 years. 

This boy was admitted to hospital on the eighth day of illness, complaining of 
‘ fever and pains in the stomach.’ His temperature was 103° F., the pulse was 
rapid, 124 per minute, the tongue coated but with red tip and edges. The abdomen 
was tumid, but liver and spleen were both palpable. P. Jalciparum was found in 
smears of blood sent up on the following day, and on the tenth day of illness a 
Widal test gave a strong agglutination of B. typhosus in high dilution. Progress 
was very satisfactory ; the temperature never rose above 103° F., and reached 
normal on the eighteenth day of disease (Chart 5). 





Chart 5 














202 


Here again, in spite of the double infection with malaria and 
typhoid, the course was unusually mild ; no history could be obtained 
of any prolonged fever or of anything of a typhoid-like nature which 
might have left residual agglutinins; the relatives denied any 
previous illness of a serious nature, and the general condition was 
that of a typhoid patient—feverish with bright eyes, painful, tumid 
abdomen, and so forth. 

8. G.L., female, aged 4 years. 

This child had been ill with ‘ fever and abdominal pains ’ for a fortnight 
before being brought to hospital. On admission the temperature was 102° F., 
the tongue coated, edges and tip clean. Blood examination on the same day 
revealed the presence of P. Jalciparum. Five days later as the temperature was 
still irregular a Widal test was asked for and the result was an agglutination of 
B. typhosus ; on this occasion no malarial parasites were detected though carefully 
sought for. On the evening of the second day after this the temperature touched 
normal, rose again next day to ioo° F. and thereafter remained normal. Con¬ 
valescence was uninterrupted (Chart 6). 

The general aspect of the patient was that of a child suffering 
from enteric fever, and the rapid defervescence is again here worthy 
of note. 


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Chart 6 


1 he suggestion may be made that we were dealing here with a 
patient suffering from a mild or from an abortive attack of enteric 
fever, and that the true, if not the only, cause of the fever 
on and after admission was the presence of the malarial parasites. 



203 


Everyone knows how anomalous are the forms of enteric fever as met 
with in children, but against the above explanation is the fact that 
the temperature continuing irregular was the reason for making a 
Widal test, and that on that occasion prolonged search failed to 
reveal the presence of any malarial parasites. 

9. A.A., male, aged 23 years. 

This patient was admitted to hospital with a temperature of 103° F., a coated 
tongue, and loose stools. The duration of the present illness could not be 
determined with accuracy, because he had suffered a few weeks earlier from influenza 
and broncho-pneumonia (during the epidemic of the so-called Spanish Influenza), 
and had not thoroughly recovered from this, though convalescing and gaining 
strength, when he again ‘ felt feverish, had pains in the limbs, and severe headache.’ 
There was no history of any previous attack of enteric fever nor of any illness 
suggestive of such. Examination of the blood was undertaken both for agglutination 
reactions and for the presence of malarial parasites. P. vivax was found, and 
the Widal test gave a well-marked agglutination with B. typhosus , negative with 
the Paratyphoids. As the chart shows (Chart 7) the temperature each morning 
for four days after admission was ioo° F., and from I to 3 degrees higher each 
evening. On the fifth day it became normal, or rather subnormal, with a rapid 
fall and remained so till the patient left hospital after a stay of four weeks. Con¬ 
valescence was gradual, but steady and uninterrupted. 



Chart 7 


It will be seen here that the temperature for the short time during 
which it remained elevated after the patient came to hospital was 
not like that of benign tertian malaria, but resembled more that of 
enteric fever in the third week; the rapid, almost critical, fall, 









204 


however, during the night of the fourth day after admission is more 
suggestive of an uncomplicated malaria brought under control. The 
headache was continuous and the general appearance very suggestive 
of enteric fever, while the course of convalescence resembled this also 
rather than an ordinary attack of malaria. 

There is, of course, the possibility that the attack of 1 influenza 
and broncho-pneumonia* had been wrongly diagnosed, and had 
been enteric fever from the start; this case would then come under 
the category of malaria developing towards the end of an attack of 
enteric fever. Against this it may be stated that the patient had 
been in hospital for his attack of ‘influenza* and under the charge 
of a medical officer who is always careful to send to the laboratory 
a specimen, and if necessary repeated specimens from any case in 
which enteric fever is suspected, or, in fact, of almost any case of 
fever at all before starting to treat the patient for malaria, as so 
many do. It is a common fault, I believe, in all tropical countries 
to treat every case of ‘ fever * as malarial, and only when quinine is 
found ineffectual are means taken to verify or refute the diagnosis. 
Here, with a laboratory on the spot, this is rarely done, the blood 
being taken in the great majority of cases for examination before 
treatment is begun. 

io. A.S., male, aged 8 ye ays. 

This patient was brought to hospital with a history of having suffered from 
‘ fever and pains in the head ’ for nine days. On admission the temperature was 
only 99 0 F., the tongue was clean, there was no vomiting and no distension of the 
abdomen. The spleen could be easily felt. The case was believed to be one of 
malaria, but in view of the history of nine days’ fever a Widal test was also carried 
out. P. falciparum was found, and the Widal reaction was positive, more marked 
in high dilutions than in lower. (I have already reported several instances of this 
which, in children out here, I find by no means uncommon). Quinine, which 
was given at once, appeared to have no effect on the temperature. Some 36 hours 
afterwards it was 104° F., next morning ioo°, and the following morning 103°. 
It then came down and remained between ioo° and 102° for ten days, then 
oscillated again for a couple of days, after which time it fell to normal and remained 
there (Chart 8). 

The course of the illness and the convalescence were both like 
those usually met with in mild cases of enteric fever, the temperature 
reaching normal on the twenty-eighth day of illness and the patient 
being sufficiently convalescent to leave hospital after a stay of 
thirty-seven days, or forty-six days after the onset of the fever. 



205 


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Chart 8 


This is very typical of the old ‘ typho-malaria/ and is inserted here 
for that reason. There is nothing otherwise worthy of special note 
except that the Widal reaction was a ‘ reversed * one. 

II. A.G., female, aged n years. 

This patient was admitted to hospital on the twenty-first day of illness having 
suffered with fever, headache, and constipation. She had a coated tongue, and 
had been vomiting, it was stated, shortly before admission. The abdomen was 
neither distended, tender, nor painful; the temperature was 103° F.; the heart’s 
action was rapid, 124 per minute, and examination of the lungs showed a few 
rhonchi at the upper parts on both sides. The blood was examined both for 
agglutination and for malarial parasites. The serum agglutinated R . typhosus in 
lower dilutions (up to I in 50), but not in 1 in 100 or over ; P. falciparum was 
present, both ring-forms and crescents in considerable numbers. Blood culture 
was negative, as one would expect at this stage of disease—the end of the third 
week—but B. typhosus was isolated from the stools. The chart shows a rapid 
fall of temperature following the administration of quinine (Chart 9). 

The question here is: Can the whole condition be ascribed to 
malaria ? The clinical state certainly supports this, and the rapid 
effect of quinine further backs it up. One point against this is not 
very strong, namely, the positive Widal reaction in comparatively 
low dilutions only. Now, there was no history of any previous 
attack of prolonged fever which might allow us to interpret the 
Widal result as a residual reaction, or to the fact of the patient being a 
carrier (the presence of bacilli in the stools would support equally 
the last idea or the fact of a recent attack). One would naturally 












2o6 


expect in a definite case of enteric fever that the reaction would be 
stronger at the end of the third week of illness, and there is no 
reason for doubting the history. Some time ago, however, I met 
two cases in which, though they were undoubted instances of typhoid 
fever, the Widal reaction was negative, although repeatedly 
tried, until the twenty-eighth and thirty-second days of disease 
respectively; while a still more striking instance occurred in my own 
case. In spite of every clinical symptom indicating typhoid fever 



Chart 9 


and repeated blood examinations, it was not until the onset of a 
second relapse and the forty-eighth day of illness that a positive 
agglutination of B. typhosus was obtained, and then only in 
1 in 50, though seven to ten days later a marked agglutination in 
high dilution was obtained, and is still, though it is now over 
six months since the fever disappeared. 

All one can say in the case of A. G., is that the patient was 
excreting typhoid bacilli in the faeces while suffering also from 
subtertian malaria, but whether the presence of the bacilli was due 
to her being a carrier in the ordinary sense of the term or a patient 
suffering with a recent attack it is not possible to say with certainty. 












207 


12. L.R., male, aged 20 years. 

The statement given by this patient on coming to hospital was that he had 
been suffering with fever and headaches for two days previously ; though he 
had not been feeling really well for three or four days prior to that. The tempera¬ 
ture on admission was 102° F., the tongue was slightly coated, but moist, the 
spleen was distinctly enlarged, and the liver was also palpable. Examination 
of the blood showed the presence of P. falciparum , while a Widal test gave an 
agglutination of B . paratyphosus a, up to 1 in 50 and of B. typhosus to 1 in 30 
only. The temperature became normal five days after admission and the case 
was regarded as probably one of Paratyphoid infection only, the agglutination of 
B. typhosus being of the nature of a group reaction. Some ten to twelve days 
later the patient felt quite well, and was grumbling at being still kept in hospital, 
but on the blood being again examined, more as a matter of curiosity to see whether 
the reaction was the same as before, it was found that agglutination of B. typhosus 
W'as very marked up to I in 500 at least, higher was not tried, while B. paratyphosus, 
a, was agglutinated up to I in 50 only as before. The stools were also examined 
and B. typhosus was isolated from them ; B. paratyphosus , if present, was not 
isolated. Smears of the blood examined the same day revealed the presence of 
P. falciparum and also P . vivax , both in large numbers, one or two in every field 
of the smear, and several corpuscles contained two and occasionally three parasites. 
The patient was told that he could not leave just then, at which he was very angry, 
stating that he felt perfectly well. The temperature being taken frequently 
during the day showed a rise at 2 p.m. to 102° that day, but with very little 
discomfort, and after this never exceeded ioo°, and on treatment with quinine 
soon became normal again, and the patient left a month after his first admission 
(Chart 10). 


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There are several points worthy of note in this case: firstly, the 
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the reaction being more marked with the latter in the early stage 


















208 


and with the former later; secondly, the coincident infection with 
P. falciparum ; thirdly, the exceptional mildness of the mixed 
infections, the temperature becoming normal early in the second 
week of illness; fourthly, the rapid recovery so that the patient 
chafed strongly at being detained more than a fortnight in hospital; 
and lastly, the large infection with subtertian and benign tertian 
parasites without any feeling of malaise. 

13. F.S., female, aged 6 years. 

This child was brought to the hospital with a history of ‘ fever, pains in the 
head and stomach 9 for six days. The temperature in the taking-in room was found 
to be 102 0 F., the tongue was coated and the abdomen distended, and the child 
was admitted. Smears taken on the day of admission showed P. falciparum , and, 
as the fever persisted, a Widal test was carried out three days later and agglutination 
of B. typhosus was obtained in high dilution. Except for considerable weakness 
and listlessness during the third week of illness nothing remarkable occurred, 
and after touching normal on the seventeenth day of disease, the temperature came 
down finally two days later and remained normaF(Chart 11). 



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Chart ii 


14. S.S., male, adult, age not known. 

This man complained of feeling 1 chilly and then hot,’ and of abdominal 
discomfort for eight to ten days. His temperature was found to be 102°; the 
blood was examined and malarial parasites were found. Two days later spots 
suggestive of typhoid rash were noticed, and the stools were also ‘ typhoid-like/ 
so the blood was sent for a Widal examination and the serum gave an agglutination 
of B. typhosus up to I in 50 distinct, partial in 1 in 100. By this time the tempera¬ 
ture was normal (see Chart 12). Except for a recurrence of the malaria a week 
later there were no further untoward symptoms. 










209 



Chart 12 


The record of temperature in this patient is that of uncomplicated 
malaria. The symptoms of enteric fever, I am informed by the 
medical attendant, were quite distinct; rose spots, tender abdomen 
with some distension, offensive ‘pea-soup* stools. This man had 
never had any previous attack of fever so far as he was aware, he 
had only come out from England a few months before and he had 
never had any prophylactic typhoid inoculation. 

As the patient had been feeling ill for some eight to ten days 
before admission and the spots were first seen six days afterwards, 
it would appear that the malarial element was the main cause of 
the early indisposition. If we take it that the rash came out on 
the seventh or even tenth day of disease (enteric), the fall of the 
temperature and the generally good condition of the patient point 
to the case being an abortive one from the enteric aspect. Of a 
certainty the malaria had no adverse effect upon the enteric fever. 

15. C.P., male, aged 28 years. 

This patient was admitted to hospital stating that he had been suffering with 
‘ fever ’ for the previous nine days; apart from ‘ feeling feverish,* he stated that 
he did ‘ not feel at all ill,’ nor did he appear so. Beyond a temperature of ioi° F., 
a somewhat coated tongue and a just palpable spleen, there was nothing objective 
made out. There was no abdominal distension or pain. He had never suffered 
from any previous attack of fever of any prolonged duration, and had always been 
very healthy. The blood was examined by smears for malarial parasites, by 
cultivation and for Widal’s agglutination reaction. The smears showed a heavy 














210 


infection with P. falciparum , often as many as eight or ten in a single field, and 
several corpuscles contained two and occasionally three parasites. The Widal 
test gave a strong agglutination of B. typhosus , negative with B. paratyphosus , 
a and b. The culture was negative as regards the blood, but from a stool the 
B . typhosus was present in large numbers and was isolated. The course of the 
illness was exceptionally mild ; the patient was most cheerful and ridiculed the idea 
of his being ill enough to be kept in bed. The temperature became normal on the 
fourteenth day of disease ; convalescence and recovery were exceptionally rapid and 
uninterrupted (Chart 13). 


mm 

nta 

Id 

IC 

IC 

□1 

71KE 

IC 

IB 

m\ 

■ 

DOS 

3302 


m 

a 

392 

3332 

ib: 

132 

331 

39 

Mil 

III 

1 

III 

II 

IN 

ill 

HI 

III 

II 


HHK 

1 

l 

111 

11 

■1 

III 

in 

II 

II 


mm 

1 

11 

in 

n 

■1 

III 

in 

II 

II 


INK 

i 

III 

m 

n 

u 

III 

11 

in 

II 


Mil 

# 

III 

111 

11 


HI 

111 

H 

H 


INIS 

i 

in 

m 

n 



n 

11 

U 


m s 

1 

11 

111 

11 

■1 

IB 

1 

in 

U 


II5E 

1 

11 

111 

11 

■1 

HI 

■ 

H 

II 


Mm 

K 

11 

in 

HI 

II 

III 

l 

II 

U 


Mil 

1 

1 

ill 

II 

■1 

III 

111 

II 

U 


MIA 

1 

IVS 

ii 

ni 

u 

IUI 

IH 

H 

II 


■Ml 

■IN 

11 

m 

IB 

MM 

m 

181 

ll 

■1 



Hill 

n 

m 

in 

III 

IIH 

mm 

II 

III 

III 

■■ 

Mil 

iiggi 

III 

III 

III 

INI 

IUI 

II 

III 

III 

■U 

■in 

uni 

m 

in 

III 

IMI 

Mil 

n 

III 

III 

■■ 


Chart 13 


Very little comment is needed. There was no history obtainable 
of any previous illness pointing to enteric, and the patient was very 
intelligent and his relatives bore out his assertion. The Widal 
reaction cannot thus be interpreted as residual from a previous 
attack. Isolation from the blood would hardly be expected as 
late as the tenth day of disease; hence the attempt, which was 
successful, to isolate the organism from the excreta. He had not 
been inoculated, in fact, none of the cases here recorded had had 
inoculation. 


The number of cases described is too small to allow of any 
generalisations, but so far as one may venture on statements based 
on so few cases, and feeling that those statements may and probably 
will have to be modified as further instances are met with, one may 
note provisionally: — 






211 


1. That the effect of dual infection with malaria and enteric 

fever does not merit any graver prognosis than the latter 
alone, and that in many, if not most, of the above the 
temperature fell early and the attack proved to be mild. 

Judging from the accounts given, it may be argued that 
the cases were mild from the enteric point of view, and that 
the quinine dealt successfully with the malarial element of 
the illness. It may be so, but, on the other hand, enteric 
fever in Jamaica, at all events in Kingston, is by no means 
a mild disease. The mortality is high, and in those who 
recover, convalescence is as a rule prolonged, and the 
debility marked; therefore it is, prima facie, hardly 
credible, at least it is extremely improbable, that all these 
cases in which malaria and enteric fever were coincident 
should be exceptionally mild. 

Further, malaria here is also a serious condition; this is 
the opinion of medical men who have practised in Jamaica 
for many years, and the laboratory records show that of the 
many specimens sent up for examination 80 per cent, of 
those revealing parasites show P. falciparum, and the 
recorded mortality from malaria alone is high. 

2. We are, therefore, driven to the conclusion that the presence 

simultaneously of two diseases, one protozoal, the other 
bacterial in nature, each of which is usually severe in 
type and effects, leads to the production of an illness of 
comparative mildness in both these respects. 

3. Sir Almroth Wright’s quotation at the beginning of this 
paper is to the effect that anti-typhoid inoculation appeared 
to reduce susceptibility to malaria. If we regard the 
attack as equivalent to inoculation, then it would appear 
(as in Cases 12 and 15) that the malarial infection, though 
a heavy one, caused very little constitutional upset, and 
that the patients experienced no malaise and wondered 
why they should stay in bed when they felt quite well. In 
the above cases, however, there are instances which are in 
favour of the corollary to this, viz., that the presence of 
malaria coincidently with enteric fever appears to modify 
greatly the severity of the latter. The point I wish to 



212 


emphasise is not quite the converse of the typhoid and 
malaria statement: that as anti-typhoid inoculation 
appears to reduce susceptibility to malaria, so malaria 
reduces susceptibility to typhoid, for I do not think this is 
the case. The majority of persons out here suffer at some 
time with malarial infection, and, as stated, four out of 
five are infected with P. falciparum ; nevertheless, enteric 
fever is a severe disease here with a comparatively high 
mortality. But when the two conditions, typhoid and 
malaria, are present together in the initial stages of the 
former, the course of each is modified, and favourably so; 
in other words, the presence of malarial parasites in the 
blood appears to have a mitigating effect on the severity 
or the course of an attack of enteric fever. 

4. There arises the question as to whether, in the tropics , 

quinine has a beneficial or any effect on the severity and 
course of enteric fever. I say expressly ' in the tropics ’ 
because in temperate climates one has had repeated proof 
of the old dicta 4 any fever which does not yield to quinine 
is not malarial *, and 4 in any case of fever of more 
than a week’s duration suspect enteric.* The effect, if 
any, is but slight, judging from a very large number of 
cases of ordinary enteric fever seen here. Of course, in a 
patient who has been subject to malarial attacks the 
exhibition of quinine may modify slightly the course of 
fever at the onset of enteric, but as a rule it is the failure of 
quinine to mitigate or reduce the temperature that brings up 
the suspicion of the presence of enteric fever, especially 
when the blood is not submitted to examination for 
malarial parasites. In other words, from experience of a 
large number of cases one may say that quinine has very 
little effect on ordinary cases of enteric fever. 

5. Can malaria of itself lead to a positive Widal reaction in the 

absence of enteric fever? (It may be stated again here 
that none of the cases quoted had ever been given anti¬ 
typhoid inoculation, so that the reaction obtained was not 
due to inoculation agglutinins.) 

With a view to testing this aspect of the question, 



213 


I have for some months past, when taking smears from 
patients suffering from malarial fever, taken blood also 
for the agglutination reaction. I have tested some two 
hundred cases in this way and in no instance have 
I obtained definite positive results in uncomplicated 
malaria. Occasionally a loss of motility and a weak 
attempt at agglutination might be noticed in dilutions up 
to I in 20, rarely possibly i in 30, but never higher than 
this. So far, therefore, it may be said that the Widal 
reaction is not given by patients suffering from uncom¬ 
plicated malaria. We may safely say that in high dilutions 
the reaction is specific, in lower, group agglutinins come into 
play, while in very low dilutions, even normal blood at 
times appears capable of giving some degree of agglutina¬ 
tion. In the above cases the majority came to hospital too 
late for blood culture to yield successful results, and one 
had to rely largely on the Widal reactiop and the isolation 
of the bacilli from the excreta. In one case, B. M., the 
patient was seen on the fourth day, and the organism was 
obtained by blood culture. 


SUMMARY 


1. Anti-typhoid inoculation, it has been reported, reduces 
susceptibility to malaria. 

2. Enteric fever in Jamaica is a severe affection with compara¬ 
tively high mortality. 

3. Malarial infection in Jamaica is also a severe condition, in a 
large majority of instances of the subtertian variety. 

4. Quinine has not any marked effect on uncomplicated cases 
of enteric fever. 

5. The serum of patients suffering from uncomplicated malarial 
fever will not give a positive reaction to Widal’s agglutination test. 

6. Coincident enteric fever and malaria (that is, when a patient 
is seen early in the attack of enteric fever and examination of the 
blood reveals the presence of malarial parasites at this early stage), 
in many instances at least is remarkably mild, in type and in 



course, and recovery is usually rapid and complete, more so than in 
the case of either affection separately. 

I desire to express my acknowledgments and thanks to Dr. T. R. 
Matthews for allowing me access to these cases, for eliciting 
histories, interviewing friends and relatives, and facilitating me in 
obtaining specimens for examination from the patients. 


REFERENCES 


Lyon (1900). Johns Hopkins Hosp. Rep . Vol. VIII, dted by Kollc and Wassermann, ' Hand- 
buch der pathogenen Mikro-orgamsmen/ Vol. Ill, p. 777. 

Manson, Sir Patrick (1917). Tropical Diseases, 6th Edition, pp. 117, 380. 

Scott, H. H. (1913). Practitioner , Nov. 

- (*915)* Enteric Fever in Kingston and its suburbs. Ann. 7 rop. Med. & Parasitol. 

Vol. IX, p. 239. 

Wright, Sir Almroth (1919). Lancet, March 29th. 



215 




THE METABOLISM OF WHITE RACES 
LIVING IN THE TROPICS 

II. THE COMPOSITION OF THE URINE 


BY 

WILLIAM JOHN YOUNG 


From the Biochemical Laboratory of the Australian 
Institute of Tropical Medicine 


(Received for publication 17 June, 1919) 


In the course of a study of the effects of a tropical climate upon 
a white working race, a number of urines have been analysed to 
ascertain whether they showed any marked variation in the 
quantities of constituents from those given in the physiological 
text-books as the average excretions for dwellers in a temperate 
climate. 

Investigations on the urine of white men living in the tropics 
have previously been carried out by a few workers only, and these 
have been done mostly with the object of inquiring whether the 
changed conditions bring about an altered mechanism for regulating 
the thermal equilibrium of the body. 

The only systematic work of any extent is that of Eijkman 
(1893), in Java, who estimated the calorific value of the food of 
nineteen Europeans, and drew up a balance sheet between the 
nitrogen of the food and that excreted in the urine and faeces. In 
the course of these experiments he analysed the daily urine of his 
subjects over a period of fourteen days. 

A few workers have recorded the results of observations upon 
themselves made during a short visit to the tropics, and have 



216 


compared these with figures obtained in Europe before or after their 
journey. Among these latter may be mentioned Schilling and 
JafF6 (Schilling (1909)), who compared the composition of the diet 
taken by themselves in Europe with that consumed during a short 
stay in West Africa; in addition they determined the nitrogen 
excreted in the urine and faeces in both places. Rancke (1900), 
Wick (1910), Glogner (1909), and a few others have also published 
the results of a few observations made on themselves. 

The results have been recorded of the examination of the urine 
of a number of American soldiers serving in the Philippines, as well 
as of natives of these islands, but these observations have been 
mainly confined to volume and specific gravity (Chamberlain 
(1911)). 

A certain amount of work has been done on the composition of 
the urine of native races in the tropics. Eijkman (1893) analysed 
the urine of a few Malays for comparison with his Europeans, 
whilst McCay (1912) has recorded observations on a large 
number of urines of Bengalis, and Aron (1909) has examined 
urines of natives in Manilla. More recently Campbell (1917) 
published the results of analyses of the daily excretion of a few 
native medical students in Singapore. In most instances it was 
found that the urines of natives showed marked differences from the 
European standards, thus, for example, the nitrogen was generally 
much lower, which may be accounted for by the native diets being 
generally poorer in protein, and richer in carbohydrate than the 
diet usually consumed by the white man. 

The results of these previous investigations will be discussed in 
reference to those recorded in Townsville in the following pages. 

In a previous publication (Young (1915)) the results were 
recorded of an investigation carried out in Townsville into the 
partition of nitrogen amongst the various nitrogenous constituents 
of the urine. It was found that the total nitrogen in the urine was 
somewhat low, but the proportions of this nitrogen in the individual 
compounds agreed approximately with those found elsewhere. 

It is a common belief that in the tropics the excessive loss of 
water through the skin causes a corresponding decrease in volume 
of the urine, and that the continued passing of concentrated urine 
is responsible for a large amount of kidney disease. In the present 



217 


investigations, therefore, the main object has been to compare the 
concentration of urine with that found in Europe, and attention has 
been confined to the volume, specific gravity, total nitrogen, 
freezing point (osmotic pressure) and certain inorganic constituents. 


METHODS 

The following methods were employed in these experiments. 
The specific gravity was determined at I5°C. with a Mohr's 
balance, the total nitrogen by Kjeldahl's method, and the sodium 
chloride by Volhard’s method, i.e. by adding a known amount of 
silver nitrate, filtering off the silver chloride, and estimating the 
silver nitrate left unchanged by titration with potassium sulpho- 
cyanide in presence of a ferric salt. The phosphate was determined 
by titration with a uranium solution. The freezing point of the 
urines was obtained in the usual manner. 

The results are put together in the accompanying Table I, all 
the figures being obtained during the hot summer months. The 
subjects were drawn from various walks in life, and comprise 
persons doing laboratory work, a medical man in practice, a 
newspaper reporter, hospital wardsmen. several men doing office 
work in the city, a shop assistant and several labourers. The 
figures in the table refer to the whole urine passed in twenty-four 
hours, the subjects living on their ordinary diet. 

As is well known, with a freely chosen diet considerable varia¬ 
tions are observed from day to day in the quantity of the various 
substances excreted in the urine, so that no great accuracy can be 
claimed for a single twenty-four hours’ sample. The following 
figures, however, were found for a number of persons of very 
different occupations, and it will be noticed that on the whole they 
vary from the usual standard in the same way, so that the average 
figures may be taken as an indication of the type of urine found in 
tropical Australia. 



218 


Tmile I. 


Subject 

Volume 

C.C. 

Specific 

Gravity 

Total 

Nitrogen 

grams. 

Cm 

Sodium 

Total 

ms 

Chloride 
per 100 

C.C. 

Freezing 
point 
degrees C. 

P. 0 , 

grams. 

i (a) 

1116 

i*oi6 

8-7 

6*14 

°*55 

1*028 


?\ 

IOOO 

1*017 

7 -« 

3*60 

0*32 

o <935 

... 

w 

(4 

5 2 5 

1*027 

8*o 

3 * 6 o 

o*68 

I*726 

... 

5*5 

1*026 

6*8 

4*75 

0*92 

1*907 

... 

*(a) 

830 

1*023 

10*2 


0*65 

••437 


w 

644 

1*023 

8*0 


o *57 

I*466 

... 

3 (?) 

885 

1*02$ 

11*0 


0*92 

1*643 


w 

814 

1*032 

14*7 


0*85 

2 *°95 

... 

4 («) 

7*8 

1*030 

10*9 


i* 19 

2 *°49 

... 

w 

753 

1*031 

ii*7 


i *37 

2*102 


5 («) 

624 

1*032 

98 

673 

1*08 

2*119 


w 

556 

1*031 


4*50 

o*81 

2*025 

... 

6 

965 

1*026 

ii*7 

12*35 

1*28 

1*869 

... 

7 

s«* 

1*028 

6*5 

6*29 

i-o8 

,.694 

... 

S 

874 

1*027 

15*0 

4*82 

o *55 

P929 

... 

9 

738 

1*031 

14*3 

5*64 

0*76 

2*105 


lo 

852 

1*024 

ii*7 

669 

o *79 

i* 7 , 4 

... 

II 

702 

1*023 

9 * 2 

5-68 

o*81 

1*675 

... 

12 

875 

1*021 

9*7 

8*85 

1*01 

1*613 

i #2 4 

*3 

593 

1*022 

6*9 

7**4 

1*22 

, * 75 2 

I- 5 7 

*4 

*390 

1*023 

15*6 

* 5*37 

1*06 

00 

o*81 

! 5 

634 

I*03I 

u*6 

5 * 4 i 

0*85 

2*259 

2*61 

16 

9°5 

1*027 

,2 *7 

8-94 

o *99 

! * 97 ° 

2*24 

>7 

920 

1*020 

8*9 

7*65 

0*83 

1*581 

2*06 

18 

688 

1*026 

10*8 

6*03 

0*88 

1*987 

i *34 

*9 

*55 

1*025 

8*7 

5-59 

0*85 

i*6ii 

2*04 

20 

375 

I*Ol8 

3 *i 

2*95 

o *79 

1*360 

o -57 

21 

7*8 

1*026 

9*6 

6*78 

0*94 

1*925 

i*86 

22 

1070 

1*024 

> 4-3 

10*38 

0*98 

... 

2*61 

l 3 

9 22 

1*027 

,2 *4 

12*76 

i' 3 2 

... 

1*76 

2 4 

782 

1*025 

(o*8 

7*38 

o *94 

... 

1*96 

2 5 

621 

1*023 

8*8 

5*00 

0*49 

• •• 

1*56 

Mean 

782 

1*0254 

10*4 

7 *o 

1*13 

l*76l 

i *73 

Maximum ... 

1487 

1*032 

17*3 

* 5*4 

i *39 

2*259 

2*6i 

Minimum ... 

375 

1*013 

3 *i 

2 *95 

0*32 

°*935 

0*57 


* 3 i *4 2J aye tl*e average* over one week in each case. 
















219 


The total volume, specific gravity, and inorganic salts 

The figures given in the text-books (Starling (1912)) as the 
average for temperate climates are:— 



European 

Townsville averages 

Volume c.c.. 

1500 

782 

Specific Gravity . 

1*014 to 1*020 

1*0254 

Freezing point . 

-0*87° to -2*71° 

-0*935° to -2*259° 

Sodium Chloride . 

15*0 grams 

7*0 grams 

Phosphate . 

2*5 grams 

1*73 grams 


When compared with these it is seen that the volume is very 
considerably lower, the average being only 782 c.c. per day. The 
gravity is correspondingly higher. The freezing points of the 
urines come in every case within the range given for temperate 
climates, the maximum being - 2 259 0 C. with a mean depression of 
176° C. 

The sodium chloride shows a very interesting difference. The 
quantity was in every case but one (No. 8) far below the European 
figures. The average amount passed was found to be only seven 
grams per day. When the actual concentration of this salt is 
considered it is seen that it is not very different from that found in 
temperate climates, namely about 1 or V 2 per 100 c.c. The figures 
actually found vary from 072 to 179 grams per 100 c.c. with an 
average of 1*13. This small amount of salt in the urine is thus 
instrumental in maintaining the freezing point about the same as 
that found in Europe. The small total quantity of chlorides 
present in the urine may be accounted for by the excessive perspira¬ 
tion, and a quantity of sodium chloride being excreted in this way. 

The amount of water and sodium chloride lost in the sweat 

The quantity of water lost in perspiration in the tropics is very 
considerable. Eijkman (1893) estimated that in his subjects, 
laboratory workers, an average of 1730 c.c. was lost per day in the 
perspiration and from the lungs. This, however, seems a small 







220 


amount for a man doing manual labour in a hot climate such as that 
of North Queensland, and where all the work is carried out by white 
people, who work mostly to the same hours as in a temperate 
climate, that is during the hottest part of the day. Hunt (1912) in 
a paper upon the effects of a dry hot climate upon the body, 
describes his experiences during a march in the Deccan in India, 
with the dry bulb temperature at 104° F., and states that none of his 
party consumed less than three gallons (13*6 litres) of water per day, 
a quantity which he considers as the minimum advisable. He states 
that even with this quantity of liquid the flow of urine was by no 
means free. Most of this water, therefore, must have passed out by 
the skin and lungs. 

In a warm moist climate, where there is little evaporation, the 
slightest exertion causes profuse sweating, and this saturates and 
clogs the clothing, and thus further interferes with free evaporation 
from the skin. This induces further excretion of sweat, a sort of 
vicious circle is set up, and the body is not able to utilise the sweat 
economically. The result of an increased consumption of water is 
an increased perspiration, which adds to the discomfort, and it is 
only when a very large amount of water is taken that any marked 
increase in the quantity of urine is observed. In spite of this 
discomfort the slightest exertion causes an increased desire for liquid, 
and a large quantity of water is consumed, and thus a corresponding 
amount of water is excreted by the skin. 

In some experiments carried out in Townsville the loss in body 
weight was observed during a brisk walk of seventy minutes at the 
rate of about four miles an hour, the thermometer at the time being 
8o° to 85° F. dry bulb, and 70° to 8o°F. wet bulb. The loss of 
weight in some subjects reached as much as 1100 grams, most of 
which would be due to loss of sweat. Moreover, in these experi¬ 
ments only the water actually evaporated was determined, the 
subjects being weighed in their clothing, which was saturated with 
sweat, so that the total loss of water must actually have been much 
greater than this quantity. 

Losses of water such as this must be quite a common occurrence 
with people doing manual labour in the tropics. In discussing this 
question with a local carpenter, the author was informed that in the 
workshop it is customary for the men—four in number—to fill in the 



221 


morning a large bucket, capable of holding nine to ten litres, with 
water, and from time to time during the day these men dip their 
pannikins into it and drink. The whole contents of the bucket are 
consumed during the day, and it generally has to be replenished 
before the day’s work is over. In addition these men will drink a 
large amount of tea and other liquids outside and at their homes. 
The great bulk of this water must be excreted in the sweat. It 
seems probable, therefore, that with manual work at least four or 
five litres of water per day must be passed through the skin. 

With four workers in the laboratory the amount of actual liquid 
taken in was found to average between three and four litres per day 
per man, whilst the average volume of urine passed was only 
817 c.c. 

In order to estimate the quantity of sodium chloride which may 
be removed from the body in the sweat, determinations of this salt 
were made in the sweat of two subjects. The sweat was collected 
in a room in which the air was practically saturated with moisture 
at 95 0 to 96° F. The body was well washed down beforehand, so 
as to remove all old sweat residues. Generally two samples 
collected at an interval of half an hour agreed well in the sodium 
chloride content, showing that under these conditions there was no 
concentration of sweat on the skin, and that the specimens repre¬ 
sented a fair sample of the sweat excreted. The sodium chloride 
was estimated by a modification of Volhard’s method. 

With one subject, upon two occasions, samples of sweat were 
found to contain o‘36 and 0 31 grams of sodium chloride per 
100 c.c. respectively. During the first experiment the subject lost 
650 grams in body weight during an hour in the hot room. The 
loss in weight due to respiratory exchange* during this time, as 


* The lots in weight due to the respiratory exchange was calculated in one experiment. 
The expired air was measured by breathing into a Zuntz meter and a sample of this air was 
analysed for oxygen and carbon dioxide. The subject breathed into the meter for five minutes 
just before going into the hot chamber, again after thirty-five minutes, and a third time after 
seventy minutes in the chamber. The volume of CO t evolved and oxygen absorbed in c.cs. 
per minute reduced to N.T.P. were :— 




CO* evolved 

O- absorbed 

Before entering ... 

.. 

197 

216 

After 35 minutes 

.. 

294 

327 

After 70 minutes 

. 

311 

375 


An average of these rates over the whole time gives as a rough estimate 19 litres of CO* 
evolved and 22 litres of Ot absorbed, i.e. a total loss of about 7 grams in weight. 




222 


well as the loss in water from the lungs with the inspired air 
saturated at 96° to 97 0 , may be neglected, so that the loss of weight 
represents loss by sweat. The sodium chloride excreted in the 
sweat during this time was thus 2 34 grams. 

Samples of sweat from a second subject were collected under the 
same conditions, and were found to contain o'22 grams of sodium 
chloride per 100 c.c. During the hour and a half which the subject 
spent in the hot chamber, he lost 2,000 grams in weight, 
corresponding thus to 4'4 grams of sodium chloride from the skin. 
In another experiment on this subject the sweat was found to contain 
o'11 grams of sodium chloride per 100 c.c. 

Hunt (1912) found that his sweat contained o'18 to O' 20 grams 
per 100 c.c., and he obtained higher figures (o'4) with other subjects. 
The quantity of sodium chloride excreted in the sweat may therefore 
be taken as from o'l to o - 4 grams per 100 c.c. A daily loss of 
sweat of several litres would, therefore, correspond to a good many 
grams of sodium chloride, and the deficiency of this salt in the 
urines under examination (7 grams instead of 15) may easily be 
accounted for by increased excretion by way of the skin. 

The fact that so much sodium chloride is excreted in the sweat 
does not mean that the kidneys are relieved of any work, but rather 
the other way, since this salt sifter passing from the blood through 
the glomeruli of the kidney must have been subsequently reabsorbed 
in the tubules of that organ. 

The phosphates 

The quantity of phosphates expressed as phosphoric acid 
averaged 173 grams per day with a maximum of 2'6i and a 
minimum of o'57. The normal figure given by the text-books is 
2'5 grams, so that the phosphate was slightly lower than the 
standard. The concentration of phosphate was, however, much 
higher than the standard, o'25 grams per 100 c.c. as against 
o'17 grams. In samples of urine collected here, it is a common 
thing for phosphates to separate out within a very short time after 
the urine has been passed, and in many cases the urine is actually 
voided in a cloudy condition, which is due to precipitation of the 
phosphates, since it clears at once upon the addition of acid. 



223 


The total nitrogen 

The urines examined in Townsville gave an average daily figure 
of 10 4 grams of nitrogen. Considerable variations were found in 
the same individuals at different times. In the following table are 
seen the daily averages of the nitrogen secreted by four persons over 
different periods: — 


Subject 

Date 

Daya 

Volume 

Specific 

Gravity 

Nitrogen 

i 

March, 1913 . 

9 

986 

1*026 

ii*7 


December, 1913 . 

8 

772 

1*030 

n*6 


February, 1914 . 

7 

1070 

1*027 

14*3 


January, 1915 . 

7 

849 

1*026 

12*4 


Average 




12*1 

2 

September, 1914 . 

■ 

922 

1*029 

12*4 


January, 1915 . 

| 

735 

1*030 

11*2 


Average 




11*8 

3 

March, 1913 . 

mm 

621 

1*026 

8*8 


January, 1915 . 

n 

946 

1*025 

7*8 

• 

March, 1917 . 

1 

667 

... 

9*2 


Average 




8*6 

4 

April, 1914. 

mm 

782 

1*025 

io*8 


January, 1917 . 

1 

737 

1*023 

9 ' 1 


Average 




10*1 


Although in these four cases the average nitrogen differed fairly 
considerably from time to time, yet in general it was markedly lower 
than the European average, the mean of all four persons being 
10 6 grams per day. 

The figures all point to the fact that the nitrogen is below the 
European standard. 

Urines collected during the cool season 

Northern Australia has a hot and a cool season, and during the 
latter (May to October) very different climatic conditions are 




























224 


observed, the wet bulb thermometer is much lower and occasional 
spells of what is felt by the inhabitants to be cool weather are 
experienced. A few urines have been collected during this season 
for comparison, and these are given in the table below, the numbers 
being the averages of urine collected for several days. 

Table II. 


No. 

Volume 

c.c. 

Specific 

Gravity 

Total 

Nitrogen 

grins. 

NaCl. 

grms. 

NaCl. 
per cent. 

1 

H 37 

1-017 

12*1 

... 


2 

1037 

1*021 

ii*7 

... 

... 

3 

1372 

I*OI2 

io*5 

... 

... 

4 

1367 

1*014 

1 x*o 

... 

... 

5 

1214 

I*Ol6 

9*5 

9-87 

o*8o 

6 

1912 

i*oi6 

, 4'5 

9*95 

0*59 

7 

1766 

1*018 

7-9 

8*24 

0*72 

8 

1441 

1*020 

9*4 

13*58 

o *97 

9 

878 

1*027 

10*2 

9* x 5 

1*07 

10 

11 5 1 

1*015 

9*7 

5*86 

00 

Tt- 

6 

Average 5-10 

1 357 ' 5 

1*0187 

10*2 

9*44 

o *77 

Same subject 






hot Season 

77 ° | 

1*0257 

9*95 

6*55 

o*86 


These figures show, a decidedly larger volume and smaller 
gravity than those obtained in the hot season, whilst the sodium 
chloride is larger in amount, but less in concentration. The urines 
of the last six subjects are averaged separately in the table for 
comparison with those of the same subjects during the hot season. 

DISCUSSION 

The records in the literature of the composition of the urine 
passed by the white man in the tropics are not very plentiful, and 
often contradictory in the results obtained. 

Eijkman (1893) found that his nineteen European subjects in 
Java over fourteen days gave the following daily averages: — 







225 


Volume 1,442 c.c., specific gravity 1*017, nitrogen 1304 grams. 
These urines, therefore, did not differ to any marked extent from 
those found in Europe as regards concentration, volume and 
nitrogen. Other workers have recorded the results of observations 
on themselves. Thus Plehn (1908) in West Africa found that his 
urine had an average daily volume of 1,075 c.c., and a specific 
gravity of 1025, whilst Glogner (1909) found the volume practically 
identical in Sumatra and Berlin. 

Neuhaus (1893), during a journey round the world in 1893, 
tested the average volume and gravity of the urine which he 
passed daily at various places during his journey. In the tropics 
the volume varied from 1,100 to 1,200 c.cs., and the specific gravity 
from 1 029 to 1*033, whilst outside the tropics the volumes ranged 
from 1,353 to 1,609 c.c., with a gravity of 1*021 to 1*023. His 
urine was thus more concentrated in the hotter parts of the world. 

Chamberlain (1911) took the gravities of single specimens of 
urine of five hundred and ninety-six soldiers in the Philippines upon 
two occasions at a year’s interval, and he obtained the figures 1*099 
and 1097. He concluded that the urine was therefore little 
different in gravity from that found in a temperate climate. 

In a recent research Campbell (1917) found that his own urine 
in Singapore conformed in volume and gravity to the European 
standard, being 1,560 c.c. and 1*012 respectively. It is noteworthy 
in the last instance that the sodium chloride content averaged only 
8*10 grams, or 0*52 per cent. 

When the averages found in Northern Australia are compared 
with the figures quoted above, it is seen that they differ in that the 
volumes obtained elsewhere were much higher than those recorded 
in Australia, whilst the specific gravities were generally lower. 

A comparison of the nitrogen with that found in Europe and by 
other workers in the tropics shows also certain differences. Our 
ideas of the metabolic changes which the protein taken in the food 
undergoes in the animal body have altered considerably in the last 
decade. The modem view assumes that the proteins are broken 
down by the digestive enzymes of the intestines into their constituent 
amino acids, which are then absorbed. The greater part of these 
are denitrified, the bulk of the nitrogen being quickly eliminated in 
the urine as urea. A small portion goes to make good the wear and 



226 


tear of the tissues. The nitrogen in the urine represents, therefore, 
mainly the substances produced directly from the proteins of the 
food (exogenous metabolism, Folin), which vary in quantity with 
the amount of food taken. In addition it contains the nitrogenous 
substances such as creatinine produced by metabolism of the tissues 
(endogenous metabolism, Folin), which, according to Folin, are not 
materially different in quantity whether the diet is rich or. poor in 
protein. The nitrogen in the urine is not an accurate measure of 
the quantity of protein metabolised, since a certain quantity is 
excreted into the intestine and passes out with the faeces, a certain 
quantity may be retained in the body, and a small amount is also 
lost in other secretions of the body such as the sweat. Still the total 
nitrogen in the urine may be taken as a rough indication of this 
protein. 

In ordinary life when an indiscriminate diet is consumed, 
naturally great Variations occur from day to day in the quantity of 
nitrogen passed in the urine, but the average figure as given in the 
text-books on physiology for temperate climates is about 15 or 
16 grams of nitrogen per twenty-four hours. 

Pfliiger and Bohland, and Bleibtreu and Bohland (quoted by 
Eijkman) give rather less than this, namely 12 67 and 14 93 grams 
per day. 

With regard to the nitrogen in the urine of white people in the 
tropics, Eijkman found that the urine of his nineteen European 
subjects on an ordinary mixed diet contained a daily average of 
1304 grams. He concluded, therefore, that an acclimatised 
European in the tropics passed as much nitrogen in his urine as he 
did in Europe. 

Schiller and Jaffe (Schilling (1909)) during a short visit to West 
Africa, carried out experiments on themselves, and compared these 
with observations made in Europe before and after their journey. 
They found that in both parts of the world nitrogen equilibrium was 
approximately maintained on 17 grams of nitrogen a day, so that no 
appreciable difference was observed in the urine nitrogen. 

Similarly Campbell (1917) in Singapore found that his own daily 
urine contained on the average 15*3 grams of nitrogen, similar again 
to the European standard. 



227 


These results would indicate that the quantity of protein required 
in the tropics is not different from that consumed elsewhere, and in 
fact Eijkman’s analyses of the food eaten by his subjects, conformed 
in every respect to the usual standards. A similar conclusion was 
also arrived at by other researchers. 

On the other hand Ranke (1900) published a comparison of his 
own diet during a visit of a few months to Brazil. This diet showed 
that although the protein was not much different, yet the total 
calorific value was much less in the tropics than in Europe. Glogner 
explained Ranke’s observations as probably due to a loss of appetite, 
which he states is a common experience during the first period of 
residence in the tropics, and he concluded that Ranke had not 
resided in the tropics long enough to have passed this stage. 

With regard to the nitrogen in the urines analysed in Tqwnsville 
the average obtained was very much lower than that of the European 
text-books, and did not agree, therefore, with the results recorded 
by others in the tropics. 

A certain difference in the nitrogen might be expected in a hot 
climate, on account of the larger quantity of water excreted by the 
skin, which takes with it a small quantity of nitrogen. Eijkman 
found the nitrogen lost in this way by Malays to be about 076 to 
1*36 grams, and he estimated that Europeans in Java lost about 
16 grams per day. 

Benedict (1906), outside the tropics, found that during rest the 
nitrogen secreted in the sweat was approximately 0 071 grams per 
day, but during muscular exercise he found as much as 013 to 
o*22 grams per hour. 

In experiments carried out in this laboratory and already 
referred to, samples of sweat were obtained and determinations were 
made of the nitrogen in them. This was found to be 0 040 and 
0*057 grams per 100 c.c. upon two occasions in one subject, and 
0*033 and 0*030 grams per 100 c.c. in the other; the two subjects 
thus lost only 0*26 and 0*66 grams of nitrogen respectively during 
an hour in the hot chamber. 

Allowing a man to lose from three to four litres of water a day 
by way of the skin, this would only account for a daily loss of 
1 to 2 grams of nitrogen by this means. If this be allowed for it 
would bring the average nitrogen to from 11 to 12 grams per day, 



228 


and a further allowance of io per cent, lost in the faeces would bring 
the total to about 13 to 13 5 grams, which, taking the higher figure, 
corresponds to about 87 grams of protein, a figure still below that 
usually accepted as a standard (viz., about 100 grams (Voit)). 

Eijkman found that his subjects in Java consumed on an average 
99*6 grams of protein, of which 88*2 grams were actually absorbed, 
the rest being lost in the faeces. Taking Eijkman’s average figures 
for nitrogen in the urine, 13 04 grams, and making the above 
allowances for sweat and loss in the faeces, it works out at about 
101 grams protein per day, a number in close agreement with that 
found. It would appear, therefore, that with the subjects experi¬ 
mented on in Townsville the protein actually katabolised was less 
than that found by Eijkman in Java. 

When an explanation is sought for these different results, a 
possible one suggests itself in the different conditions under which 
life is carried on in the Australian tropics. Eijkman's subjects, for 
example, were living in Java, where cheap native labour is to be 
obtained in abundance, and it is the rule to rest during the early 
and hot portion of the afternoon. In Queensland, on the other 
hand, as has been pointed out already, work is carried on by white 
people, and it is the exception to rest during the hotter hours of 
the day. 

The results of examinations of the urine of native races in the 
tropics show differences from that of white people. As a rule, the 
native consumes less protein and more carbohydrate than the white 
man, and as might be expected, therefore, the urines are lower in 
nitrogen. Eijkman found that the daily urine of thirteen Malays 
gave an average of only 8*08 grams of nitrogen. Campbell, for 
native medical students in Singapore, obtained figures varying from 
6*64 to 9*25 grams of nitrogen per day. On the other hand, Aron, 
in the Philippines, found a higher number, the nitrogen in the daily 
urine of the Philippino being stated to vary from 10 to 12 grams. 

McCay (1912) has made a large number of observations upon the 
urine of Bengalis in India, and has compared them with the 
standards for Europeans. Very decided differences were found to 
exist, as is seen in McCay’s table which is given below. However, 
if the Townsville urines be placed side by side with this table, the 
differences from the European standards are almost as striking for 



229 


some constituents as those given for the Bengalis. The urea and 
sulphate given below were obtained from five people, and are the 
daily averages for a week in each case. 

Table III 



European 

Bengali 

Townsville 

Volume c c. . 

1440 

1200 

784 

Specific gravity . 

IC20 

1013 

1025 

Urea grams . 

35 

13 

20*7 

Total nitrogen grams ... 

t8 

6 

io*4 

Freezing point . 

-2-5° C. 

-1*24° C. 

-1*76° C. 

Chlorides grams 

15*00 

10*00 

7*53 

Phosphates grams. 

3*50 

o*q8 

i *73 

Uric acid grams ... 

o *75 

o *45 

0*48 

Sulphates grams.j 

| 2*50 

i*88 

2*01 


The average daily excretion of nitrogen found in Townsville 
(10*4 grams) may be compared with that obtained by Chittenden 
(iq 11) from one hundred and eight university students in the 
United States, the average daily nitrogen in the urine being 
12*87 grams. 

It is interesting also to compare the samples for one week obtained 
from four men doing laboratory work in Townsville with those 
averages given by Hamill and Schryver (1906) for seven men doing 
similar work in the laboratory of University College, London. 



Weight 

kilos 

Total N. 
grams 

I 

67 

143 

2 

6t 

12*4 

3 

6? 

io*8 

4 

48 

8*8 

Average 

59-8 

1 1*6 

Schryver and Hamill 

72-4 

! 3*5 






23° 


Although the average nitrogen is actually less in Townsville, yet 
when considered per kilogram of body weight the figures are 
practically the same (0194 and 0186 grams), and both correspond 
to an amount of protein very much below what used to be considered 
as the daily standard (approximately 100 grams of protein), even 
when all due allowances are made for the nitrogen lost in the faeces. 


ALBUMINURIA 

It has frequently been stated that the higher concentration of the 
urine in the tropics causes a greater amount of kidney disease than 
is found in temperate climates; moreover, in the opinion of many 
medical men in Northern Australia, a greater incidence of kidney 
troubles is found there. As reliable figures to test this could 
not be found, an attempt was made to obtain some indication by 
ascertaining the prevalence of albuminuria in Townsville. 

A number of samples of urines from out-patients of the General 
Hospital were collected and tested for the presence of albumin. In 
no case did the subject show any other clinical symptoms of kidney 
trouble. The tests employed were the boiling test, the salicyl 
sulphonic test and the ferrocyanide test, and no urine that did not 
give all these tests was accepted as definitely containing albumin. 
The urine from six hundred and sixty-three patients was examined, 
including three hundred and sixty men and three hundred and three 
women, and a positive reaction was obtained in fifty-seven cases, 
forty-two of them being men and fifteen women. This would 
correspond to a total percentage of 8*6; amongst the men the 
percentage was 11 *7, and amongst the women 4*9. 

No attempt is made to draw any conclusions from these figures, 
but it is thought of interest to put them on record. The much higher 
percentage amongst men than women is worthy of note, nineteen out 
of forty-two of the cases in males being in men above the age of 
forty years. 



231 


SUMMARY 

The twenty-four hours* urine collected from a number of persons 
living in North Queensland and of different occupations was 
analysed. The daily volume was very much less than the European 
standards given in the text-books, the average volume being only 
784 c.c. This volume was increased considerably in the cooler 
weather. 

The specific gravity was very much higher, whilst the freezing 
point did not differ very much from that, found in Europe, thus the 
osmotic pressure was not very much higher. 

A striking difference was noticed in the quantity of sodium 
chloride excreted in the urine, which was very low, and this may be 
accounted for by the large loss of water in the sweat which carries 
with it this salt. It is calculated that a man doing manual labour 
in the tropics must lose several grams of sodium chloride per day 
through the skin, which would readily account for the deficiency in 
the urine. 

The total nitrogen showed a lower figure than that found in 
Europe, which cannot be accounted for by loss of nitrogen from the 
skin, since it is shown that this can only amount to 1 or 2 grams per 
day under normal circumstances. 

These results differ from those obtained by Eijkman and other 
observers in other parts of the tropics. 

An examination of the urine for albumin of persons not showing 
any other symptoms of kidney disease showed a high percentage of 
albuminuria, which was more marked in men than in women. 



REFERENCES 


Aron (1909). Philip, Journ. Sci. B. Vol. IV, p. 225. 

Benedict (1906). Journ. Biol. Cbrm. Vol. I. p. 263. 

Campbell (1907). Journ. Straits Branch R. A. Soc ., p. 57. 

Chamberlain (1911). Philip. Journ. Sci. B. Vol. VI, p. 427. 

Chittenden (1911). Quoted by McCay (1912), p. 118. 

Eijkman (1893). Virchow's Archtx. Vol. CXXXI, p. 170, and Vol. CXXXIIl, p. 105. 
Clooner (1909). Arch. Schiffs. Trop. Hyg. Vol. XIII, p. 169. 

Hamill and Schryver (1906). Proc. Physiol. Soc. in Journ. Physiol. Vol. XXXIV, p. 10. 
Hunt (1912). Journ. Hyg. Vol. XII, p. 482. 

McCay (1912). The Protein Element in Nutrition. London, p. 156. 

Neuhaus (1893). Virchow's Arch. Vol. CXXXIV, p. 373. 

Plehn (1908). Quoted by Wick (1910). 

Ranke (1900). Quoted by Schilling (1909). 

Schilling (1909). Tropenhygiene. Leiprig, p. 511. 

Starling (1912), Principles of Human Phyriologj, London. 

Wick (1910). Arch. Scbijfs. Trap. Hyg. Vol. XIV, p. 106. 

Young (1915). Ann. Trop. Med. & Parisitol. Vol. IX, p. 91. 



m 


ON THE ENDEMIC TSUTSUGAMUSHI 
DISEASE OF FORMOSA 

BY 

JURO HATORI 

MEDICAL INSPECTOR) GOVERNMENT OF FORMOSA 


{Received jor publication May , 1919) 

Plates VIII and IX 

I. INTRODUCTION 

In the Karenko District of eastern Formosa there is endemic in 
certain localities an exanthematous fever. Attention was first called 
to it in 1908 by its prevalence among the police engaged in building 
guard-lines against the savage Batran tribe of the Mokkui valley. 
Hence the disease goes by the name of 4 Batran or Mokkui fever/ and 
is also called ‘ Horin fever,* as it frequently affects people who enter 
the virgin forests adjoining the village of Horin. Further, of recent 
years the fever has appeared among inlander* immigrants in the 
Yoshino, Toyoda, Hayashida and other plantations, where a number 
of people fell victims to the disease. 

During the summer campaign of 1914 against the Taroko head¬ 
hunters of East Formosa, while acting as chief Medical Officer, 
I was able to examine minutely cases of the fever occurring chiefly 
in the Mokkui valley. My observations made it clear that the 
fever, which was always accompanied by swelling of the lymphatic 
gland system was analogous to the Tsutsugamushi or Kedani 
disease well known in the northern districts of Japan proper. 
I afterwards made more detailed investigations and laboratory 
experiments by permission of the Chief of the 1 Commission devoted 
to the Study of the Endemic and Epidemic Diseases in Formosa/ 
and the essential features of my five reports on the disease from 1914 
to the present time are summarized in the present paper, in which 
I use the familiar term * Tsutsugamushi disease/ though originally, 

* By the term ‘ inlander * the Japanese settlers in Formosa are meant. 



234 

from the clinical point of view, I called the disease ‘ exanthematous 
bubonic fever.’ 

II. BN DEM IOLOG1C AL 

A. Geographical Distribution 

The localities where the fever prevails or mite-infested areas 
exist, or are supposed to exist, are as follows: — 

(a) The Karenko district. This district is more intensely 
infected than others, the following foci being known:—(i) Mokkui 
and Rokei valley, (2) a part of Yoshino plantation, (3) Kotobuki- 
mura and the western part of Toyoda plantation, (4) Hayashida 
plantation, Horin-sho and neighbouring forests, (5) Maribashi 
plantation, (6) Seisui valley, (7) Shin jo and Takkiri valley, etc. 
Some details of the prevalence in these localities are given later. 

( b ) The Giran district. In 1914, during the operations against 
the Nanwo tribe of savages, a certain number of policemen were 
attacked by the disease while serving in the southern branch 
of the Dainanwo river valley. Five typical cases came under my 
observation. In 1918, the total number of patients (including out¬ 
patients) infected in the Dainanwo valley amounted to about fifty, 
although cases had not been reported in the preceding three years. 

(c) The Toyen district. In 1918 (November 15th), a police- 
inspector, stationed at the aboriginal boundary line of Mount 
Kappan, was admitted into the Government Taihoku Hospital 
suffering from a typical attack. It is evident, therefore, that an 
endemic focus exists in the mountainous areas. 

(d) The Shinchiku district. In 1917, at least one typical case 
was recorded among the police stationed in the village of Shakaro. 

(e) The Kagi district. Several cases of the fever were recorded 
by Drs. Kato, Sano and Senouye in 1911-13. They were said to 
have been infected in the forest region of Mount Ari. Recently a 
case occurred in the savage village of Tappan. So that it is clear 
that the disease occurs in mountainous areas. 

(/) The Ako district. In 1914, three policemen, two in an 
aboriginal village of the Airyo valley and one at Maruyama, near 
Shirinkaku, Koshun, were affected by the fever. In 1915, one police 
sub-inspector. In 1916, two Japanese and three Formosans. In 
1918, two Formosans were affected in the plains from Ako to 
Choshu. 



Ehdemie locality-Aborigine boundary 

. District boundary 











236 


From other districts of Formosa, viz., Taihoku, Taichu, Nanto, 
Tainan and Taito, no cases have been reported, and no mite areas 
are at present known. 

B. Details of the Prevalence in the Karenko District 

(a) The expeditionary police forces. Twenty-five cases, of which 
four proved fatal, were reported among the Batran sub-division of 
three thousand eight hundred men during an eighty days' campaign. 
Another sub-division of one thousand two hundred in the Takkiri 
valley had only four cases, one of which ended fatally. Not a single 
case was reported throughout the whole campaign among the military 
division which manoeuvred across the central mountain range 
running from the west to the east coast. 

(1 b ) Boundary line Guard. Among the Japanese police serving 
on the guard line between Hoppo and Keiko, a thirty-mile front, 
twenty cases were reported during 1913-14, two of which proved 
fatal. Of the guard posts on the line, those along the Rokei 
river and Rigyo lake, and also those on the Tamoran hill, were so 
seriously affected with the disease that nearly all the guards and 
their families fell sick one after another. 

(c) Inlanders' plantations. Three plantations of immigrants from 
Japan, i.e., the Yoshino, Toyoda and Hayashida plantations, were 
established under the control of the Government Industry Bureau 
in the District. The Yoshino plantation, founded in 1911, lies close 
to the foot of the mountain range, two miles westward of Karenko 
town. Fever cases appeared among the immigrants in 1912. It is 
said that it is at the foot of the mountains on the site of deserted 
aboriginal villages that people get bitten by some unseen insects. 

On the Toyoda plantation in 1914, thirty-eight immigrants 
suddenly fell sick after having been engaged in felling a forest west 
of the railway. 

In 1915, a colony of fifteen houses was established in this area, 
so the fever still persists. 

The Hayashida plantation, situated a few miles eastward of the 
Horin village, was established in 1914. In the first year the fever 
raged so violently among the new settlers that thirteen out of ninety 
cases of the disease proved fatal; subsequently the epidemic declined 
year by year. 



2 37 


The morbidity and mortality in these three plantations are 
shown below. 


Table I. 


Tsutsugamushi disease in the three plantations. 



Yoshino 

Toyoda 

Havashida 

Total 


Cases 

Deaths 

Cases 

Deaths 

Cases 

Deaths 

Cases 

Deaths 

*9*3 . 

25 

2 

0 

! 

O 



1 

2 5 

2 

*9*4 . 

3 ° 

t 

3* 

O 

90 

*3 

158 

*5 

*9*5 . 

27 

4 

32 

I 

49 

6 

108 

11 

1916 

7 

1 

0 

3o 

O 

10 

2 

47 

2 

* 9*7 . 

1 ■■ 

3 

8 

O 

3 

! 

1 

23 

4 


( d ) The Horin-sho and Maribashi plantations . Horin and the 
neighbouring forest region have been noted for the disease, as the 
latter bears the name ‘Horin fever’ in the district. In 1910, 
members of an engineering party of the Government railway depart¬ 
ment fell sick with the Horin fever. Cases also occurred among people 
working in the forest, such as camphor collectors, wood choppers, 
charcoal burners; and even farmers are also very often affected 
with the disease. When the Maribashi plantation of the Yensuiko 
Sugar Manufacturing Co., consisting of inlanders and Formosans, 
was founded in 1914, there were twenty-five cases, with two deaths, 
but in the following years the fever declined. 

( e ) Other localities . There are cases notified from other localities 
of the district, chiefly among camphor collectors and guaidmen on 
the Seisui valley, tunnel workers at Sappa, sugar-cane planters in 
Kada and Kotobuki-mura, Getsubi and Kompo-sho, etc. 

The following table gives the figures of cases notified in the 
Karenko district during 1913-17 for Formosans and Japanese: — 


Table II. 


Tsutsugamushi disease in the Karenko district during 1913—17. 



Cases 

Deaths 

Percentage 

*9*3 . 

60 

4 

6-6 

*9*4 . 

305 

33 

io«8 

*9*5 . 

118 

11 

9*3 

1916 . 

60 

5 

8-3 

1917 ••• _. 

72 

6 

8-3 







238 

The endemiological conditions in Karenko district are as 
follows:— 

(1) Local features. Noxious, i.e., mite-infected areas, are always 
limited (1) to. the. valleys of rivers; (2) to uncultivated but fertile 
fields covered with tall grass, mostly lying at the foot of mountains; 
and (3) to the flat or sloping ground covered with thick forest. 

(2) Influence of seasons. The fever generally makes its 
appearance in April and persists till July, then it declines for a 
while, only to rise once more in October, subsiding in November, as 
the following curve shows: — 



Monthly occurrence of the Tsuttugamushi disease in Yoshino and Hayashida plantation, 

1914-15. 






































239 


(3) Race . Inlanders, especially fresh immigrants, show the 
greatest susceptibility toward the virus, while the Formosans are far 
less susceptible and the aborigines seem to be almost immune against 
the infection, although one old aboriginal informed me that * patau,' 
the minute red mite, which is so abundant in the noxious areas, 
caused fever by its bite. 

(4) Age and sex. Adult men, being much more exposed to the 
danger of being bitten by the red mites than old men, children or 
females, show a correspondingly higher case incidence. 

Table III. 


Shewing influence of race and sex. 



> 9*4 

1 9 1 5 

Cases 

Deaths 

Percentage 

Cases 

Deaths 

Percentage 

Japanese male . 

242 

27 

IP15 

77 

7 

9*°9 

female 

57 

4 

7*00 

23 

2 

8-84 

Formosant male 

*4 

2 

142 

*7 

2 

11 77 

female 

1 

0 

... 

1 

O 

... 


(5) The annual case incidence and mortality. These rates vary 
greatly according to locality and year, as shown in the following 
table: — 

Tabli iv. 


Caic-incidence and mortality in the three plantations. 


Plantation 

Population 

Cases 

Case 

incidence 
per 1,000 

Deaths 

Mortality, 
per 1,000 


Yoshino. 

1,236 

27 

218 

1 

0*81 

* 9 * 4 ‘ 

Hayashida 

522 

89 

170-3 

*3 

24-9 


L Toyoda. 

681 

38 

557 

0 

... 


f Yoshino. 

1,604 

2 7 

16*83 

4 

2-49 

* 9*5 

Hayashida 

634 

49 

77-30 

6 

9-46 


{Toyoda. 

73* 

32 

35**3 

1 

**35 


Yoshino. 

1,694 

7 

4**3 

0 


1916' 

Hayashida 

729 

10 

* 37 * 

2 

274 


Toyoda . 

863 

30 

3475 

0 



Yoshino. 

1,823 

12 

6-6 

3 

1-64 

1917' 

Hayashida 

740 

3 

4-05 

1 

*35 


Tovodn ... 

1,100 

8 

°73 

! 0 










240 


III. ETIOLOGICAL RESEARCHES 

A. Preliminary Examinations on the Causative Virus 
and Mode of its Invasion 

In 1914, I made a series of bacteriological investigations in order 
to determine the causative organism of the so-called * undetermined 
fever* of the Karenko lazaret, and came to the following results: — 

(a) Serum reactions with typhoid and paratyphoid bacilli, and 

with Horiuchi's bacillus of the Manchurian exanthematous 
fever, proved negative. 

(b) All attempts to cultivate a specific organism from the 
patients* blood, urine and dejecta as well as from the 
spleen, post-mortem, failed. 

( c ) Smear preparations were made from patients* blood, 

lymphatic glands, and cutaneous ulcers, as well as from the 
spleen and liver, post-mortem, and stained with Giemsa. 
Neither bacterial nor protozoal organisms, nor cell inclu¬ 
sions could be found. 

( d) Finally, blood drawn from the vena median1 of a patient at 
the acme of the fever was injected hypodermically into a 
native monkey (Macacus cyclops) with a definite positive 
reaction. 

There is, on the other hand, no doubt about the fact that the 
virus invades the body through the skin, because, when carefully 
examined, an ulcer, the seat of invasion, is to be found in all cases, 
and it is recognizable where the mite has attacked the patient. 
It is usually accompanied by swelling of the adjacent lymphatic 
glands, or in other cases by a lymphangeitis apparently connecting 
the spot with the lymphatic gland. From these facts I easily drew 
an analogy between this endemic fever and the Tsutsugamushi 
disease endemic in the northern part of Japan, caused by the bite 
of the tsutsugamushi or akamushi mite. To ascertain the accuracy 
of the inference, I collected and examined rats in an endemic area, 
and found, as I expected, red mites parasitic on the rodents, and 
these are in every respect identical with the Japanese examples. 

My preliminary conclusions concerning the etiology of the 
endemic fever are as follows: — 

(1) The causative virus of the endemic fever is obscure, very 
likely of an ultra-microscopic nature. 



H 1 


(2) Invasion, without exception, takes place through the 
epidermis. 

(3) T his is effected through the bites of the minute red mites 
found in the endemic areas. 

B. The Tsutsugamushi or Akamushi, the Virus 

Carrier. 

Zoological studies of the Japanese red mite, the fever 
transmitter, well known as tsutsugamushi, akamushi or kedani, 
have been reported, each independently, by Profs. Nagayo, 
Kawamura, Miyajima, Tanaka, and their collaborators, and the 
life cycle of the mite has now been made clear by the discovery of 
its parental form by these authors in 1916. According to my 
researches, the Formosan species of the pathogenic red mite is 
identical with the Japanese tsutsugamushi or akamushi, the larva of 
Trontbicula akamushi * Brumpt. And the parental forms of our 
species agree with those of the Japanese ones. 

In the endemic area in Formosa, the red mites are found in 
nature parasitic on such rodents as Mus rattus rufescens (the 
common house rat of the island), M. dectimanus , M. musculus , 
M. agrarius , etc., or on such insectivora as Crocidura muschata , 
commonly swarming in the interior of the ear of the animals. 
Sometimes pheasants, chickens, even dogs and cats, are infested by 
the mites. People coming out of a mite-infested field or forest 
occasionally carry a number of mites attached to various parts of 
the body such as the groin, scrotum, axilla, etc. 

The parasitic red mite, ‘patau/ of Formosan Ami aborigines, is 
the hexapodal larva of Tr. akamushi , Brumpt, of ovoid shape and 
of orange red colour; its capitulum consisting chiefly of the buccal 
organs. The mandibles are chelate chelicerae, each terminating in 
a single hook. The palpus is stout, consisting of five segments, of 
which the last one bears long hairs, and the penultimate one a claw. 
The scutum on the anterior part of the dorsal surface of the body 
is somewhat rectangular, and bears seven plumose hairs. One pair 
of the later is long and tactile (pilus tactilis ) and attached to the 
pseudostigmata. One pair of eyes, most conspicuously red in 
colour, each consisting of two groups of four to five granules, is 

• Given as Leptus akamushi (Brumpt) Miyajima and Okumura. Kitasata Archives. 

Vol. I, No. 1, p. 8 (19*7)- Eds. 



242 


placed on the dorsum near the posterior comer of the scutum. The 
integument is soft and finely folded, bearing a certain number of 
plumose hairs (about thirty pairs), which are curved, with branched 
lateral ones mostly on the convex side. The type of our mite is the 
so-called ‘thin haired/ owing to its sparsely hirsute character in 
contradistinction to the so-called ‘ coarse haired * species, Tr . pseudo - 
akamushi , Tanaka, another Japanese species. The legs are strong 
and have five segments excluding the coxal one. Each terminal 
segment carries three prominent claws. On the coxal segment of 
the anterior legs the stigmatic spiracle is distinctly seen. The 
following are the measurements of the akamushi from the Karenko 
district: — 



Free in Field 

From a Rat 

From a Man 


m.m. 

m.m. 

m.m. 

Body length . 

0*196 

0*265 

o *4 

Body width . 

0*16 

0*2 

0*3 

Capitulum, length . 

0.085 

0*63 

0*09 

Capitulum, width. 

0*057 

0*05 

005 

Scutum, length . 

0*066 

0*066 


Sensory hair, length . 

0*044 

0*04 

... 

Distance of spiracles . 

0*096 

0*104 

0*127 

Distance of eyes. 

0*072 

0*074 

0*085 

Palp, length . 

... 

0*052 

0*054 

Palp, width . 

j 

0*022 

0*022 

Chclicera hook, length . 

0*022 

0*022 

OC27 

Body hair, length. 

0*057 

0*048 

0*05 

Leg I, length including claw 

0*204 

o*io 

0*21 

Leg II, length . 

0*16 

0*156 

0*165 

Leg III, length . 

0*22 

1 ; 

0*2 

0*23 


The akamushi is active in its movement, and attaches itself to 
the host with alacrity. While it sucks, it continually moves its hind 
legs; and when filled to repletion it becomes paler in colour, oblong 
in shape and markedly larger. When an ablated rat's ear, with 











2 +3 


numerous mites, is kept in vitro with soil, the mites leave the host, 
and the fully fed ones creep into the soil where they moult and 
become nymphs. As observed in deutovum, the puparium has a 
peculiar spur on the anterior dorsum. After several days of the 
resting stage an eight-legged nymph emerges. The nymph is of 
the peculiar form of a figure of 8, characteristic of the genus 
Trombicula of Berlese, the abdominal constriction dividing the body 
into two distinct parts. The cephalothorax bears the mouth parts 
and the anterior pairs of legs, while the anterior part of the 
abdominal region carries two pairs of posterior legs. The colour of 
the body is pale, faintly reddish, and densely covered with 
colourless hairs. The mandibles are chelate, and the palpi consist 
of five segments; the terminal segment is calvate, while the 
penultimate has a single claw-like process. On the area sensilligera 
of the crista metopica, a pair of long tactile hairs (pili sensoriales) 
occur. The eye is of rudimentary character, has no lens and is 
represented by a mass of reddish corpuscles. On the ventral surface 
of the distal portion of the abdomen are the genital orifice with two 
pairs of suckers, and the anus. The leg consists of six joints, 
excluding the coxal joint, end tarsi bearing two claws, with no 
pulvilli. The first pair of legs is the largest of all, its apical joint 
being the strongest. A specimen of nymph bred in vitro gave the 
following measurements: — 


Body length (with hairs) . 

m.m. 

. 0*65 

Cephalothorax, length . 

. 0-26 

Abdomen, length . 

. o *45 

Abdomen, width . 

VO 

6 

Mandible, length . 

. 0*2 

Palp, length . 

. o-i6 

Genital orifice, length. 

. 0*0$ 

Leg I, length . 

. °-4 

Leg II, length. 

. 024 

Leg III, length . 

. °-23 

Leg IV, length . 

. 0*3 

Posterior body hair, length . 

. 0-047 

Sensory hair, length . 

. 0-13 

Terminal Joint of Leg I . 

. O-II X 0-044 


An attempt to develop the nymph on vegetable matter in vitro 
was unsuccessful. At the end of the nymphal stage it passed again 



















244 


into a 
mite. 


quiescent condition and in due time moulted into the adul 


The adult form of tsutsugamushi or akamushi is found in the 
soil of infested fields, it is a tiny creature of the peculiar 8-shaped 
form; pale greyish or reddish in colour, with one pair of rudimentary 


eyes. 


The measurements of an adult male animal are as follows: — 


m.m. 

Body length . 0-94 

Body width . 0-5 

Cephalothorax, length . 0*2 

Abdomen, length . 074 

Abdomen, width . 0*5 

Palp, length . . . 0-2 

Genital orifice, length ... . 0 07 

Leg I, length . 0*62 

Leg II, length. 0*32 

Leg III, length 0*31 

Leg IV, length . 0*4 

Hair of posterior body, length . 0*054 

Sensory hair, length . o-13 

Distance of pscudostigmata . o*°54 

Terminal joint of Leg I . 0*18 x 0*07 


In a certain infested field at Karenko another species of 
Trombicula larva was found abundantly and living freely on grasses, 
and according to my observations not parasitic on man. It is 
dark red in colour, and much larger and more rapid in its move¬ 
ments than the parasitic akamushi. The adult form of this 
species is found in the soil; its body is 8-shaped, of red colour, and 
the eye is provided with a well developed lens. I provisionally 
named it Trombicula pseudo-akamushi (non Tanaka), as I am 
inclined to consider it a new species, although Prof. Miyajima 
believes that the said larva is identical with the European Leptus 
autumnalis , and that the adult animal is identical with Trombicula 
mediocris , Berl, of Java. 

















245 

Recently I found another species of red mite which infests the 
domestic fowl at Karenko. As its specific peculiarity, the fowl mite 
has fan-shaped sensory hairs on the scutum, and it is easily 
distinguished from akamushi which also often occurs on fowls. 

It seems highly probable that the mite acquires the virus from 
the adult form and transfers it to its offspring, as in the case 
of the Texas fever tick. As to the fact that rodents carry the virus, 
we can accept the authority of Profs. Miyajima and Kawamuj?a, 
who independently made successful experiments. Accordingly, the 
persistence of the disease in a definite area is readily explained. 
The further spreading of mites seems to be chiefly due to the 
migration of such hosts as rodents, birds, etc. 

In the well-known Chinese work, entitled * Honzo komoku * 
(System of Natural History), edited by Li Shiting in the 16th 
century, a certain 1 sand mite 1 has been described as a fever carrier. 
The passage reads : — 

1 In the Moorish region in South China the so-called 
“ Shashutsu ” (literally sand mite) are found in enormous 
numbers. They are so small that they cannot be seen by the 
naked eye. If people wade through shallow water or pass 
through a wood by night, the sand mite will fasten upon and 
subsequently burrow under the skin, causing a feeling similar 
to that of a slight prick, each spot being inflamed afterwards 
to the size of a lentil. The patient dies if the mite reaches 
his heart. The disease appeared first in Reinan (South 
Fookiang). In order to avoid infection, the mite on the skin 
must be scratched off with a blade of grass and some lettuce 
juice be immediately applied on the spot. If the mite lies 
already deep under the skin, it is necessary to dig it out by 
means of a needle. The mite looks like a scrabble mite.' 
(Cited from Chflgoho.) 

The sand mite of South China and the tsutsugamushi of Japan 
were first identified about ninety years ago by Genkei Ohtomo and 
his son, both experienced physicians of the Akita district. 

Now, from a zoo-geographical standpoint, Formosa belongs to 
the Indian or Oriental region, as is amply exemplified by the 
similarity of fauna, especially the venomous snakes and mosquitos. 
Therefore, it might safely be conjectured that the sand mite of South 



246 


China is probably akin to our red mite in Formosa, just as the old 
physicians deduced its identity with the Japanese mite. 

According to Dr. Schiiffner’s paper, the pseudotyphus of Deli, 
Sumatra, is caused by the bite of certain acarine larvae. Further 
South, the Mossman fever of North Australia seems to be a disease 
with a similar cause. So it would be a matter of profound interest 
to carry out further investigations on these fever-carrying mites 
distributed so widely from Northern Japan to the Far South. 
I believe that further study will very likely show a wider distribution 
of the tsutsugamushi or allied acarine mites and diseases caused by 
them in various parts of the tropics. 

C. Animal Experiments 

(a) Inoculation of the virus . Several monkeys, inoculated with 
c5 to 1 c.c. of blood from cases in the exanthematous stage, fell sick 
with fever and bubo, apparently indicating actual infection; in one 
case ( 9 ) a typical spleen was found at the autopsy. Other monkeys 
inoculated with patients’ blood or emulsion of spleen and lymphatic 
glands of infected monkeys as well as the spleen from human autopsy 
material, failed to show any signs of positive infection. 

( b ) Mite bites . Several monkeys were kept in the mite-infested 
area of Yoshino plantation. The mites were found in large 
numbers on the chest, eyelid, etc., of the animals. The bites 
were inflamed for some days, healing without ulcers. In the 
meantime the lymphatic glands became swollen and tender, and 
fever appeared after several days of incubation. The animals thus 
infected eventually recovered. 

Some species of indigenous monkeys are therefore definitely 
susceptible to the virus through inoculation of patients* blood or 
through bites of the red mite in an infected field, whereas others 
seem resjstent; this is probably due to an immunity arising from 
mite bite in their previous habitat. 

III. CLINICAL OBSERVATIONS 

A. Symptomatology 

According to my observations, the Formosan tsutsugamushi 
disease has, after the bite of infected larvae of Tr. akamushi , an 
incubation period ranging from four to ten days or more. As 



247 


prodromal symptoms, some patients complain of headache, general 
malaise, pains in the joints and loss of appetite for a few days. 

With or without these initial symptoms the fever begins with or 
without, usually with, a chill, seldom with a definite rigor. The 
body temperature gradually or rapidly rises to 39 to 4O 0 C., and 
remains at this point for a variable time, falling gradually or 
somewhat critically. The duration of the fever varies, according 
to the severity of the case, from ten days to three weeks or more. In 
grave cases, the high temperature continues for two to three weeks, 
and then lysis follows. 

In moderate cases, again, the duration is shorter, and the fever 
ends by lysis or crisis in the course of about two weeks. In still 
milder cases, the fever, generally remittent, lasts for only one to 
two weeks. Sometimes it is so slight that these cases are regarded 
as apyrexial forms. 

At the site of the bite there is a red, non-itching area, with a 
red mite in the centre, which is inclined to necrose but soon heals, 
or there may develop an ulcer, with a scab, about 5 mm. in diameter. 
The ulcer heals in one to two weeks, generally leaving some 
pigmentation. They usually occur on soft folded areas of skin, such 
as the pudenda, inner thigh, belly, scrotum or axillary region. 
The ulcer is usually single, exceptionally there are two or three. 

An important sign is adenitis in the region of the ulcer, seldom 
accompanied by lymphangeitis. The primary bubo affects the 
inguinal, femoral, axillary or other lymphatic glands, while later 
secondary enlargement of the glands occurs. The swollen glands 
vary in size, from that of a lentil to that of a pigeon's egg; they 
are single or multiple, painful on palpation. The bubo does not 
adhere to the surrounding tissues and gradually returns to its 
normal size; in one case suppuration of a cubital bubo has been 
recorded. 

Some days after the beginning of the attack, eruptions appear on 
different parts of the body, e.g., the chest, back, face and 
extremities. They are non-itching, reddish papules, and fade on 
pressure. They develop for a few days and then gradually 
disappear, leaving pigmentation. 

In grave cases, however, the papules sometimes become confluent 
and persist, mostly on the face; in mild cases they very often fail to 
appear. Icterus has never been observed. 



248 


In the majority of cases the face is congested and somewhat 
oedematous during the exanthematous period, and the conjunctiva 
is injected. In two cases I observed transitory dimness of vision 
probably due to keratitis. 

The tongue is coated and moist, but in severe cases dry. The 
pharynx and tonsils are injected and generally swollen. Nausea or 
vomiting occurs, and in grave cases hiccough. Constipation is the 
rule, but diarrhoea may occur in grave cases. 

The urine is albuminous, and the diazo-reaction is usually more 
or less positive. The spleen and liver are often enlarged and 
tender. 

Bronchitis occurred in 34 per cent, of 118 cases. The pulse is 
generally rapid, but in some cases it is slow and dicrotic, as in 
typhoid fever. In grave cases it is soft and frequent, cardiac failure 
occurring in the second or third week. Leucopenia is characteristic 
of the fever, the leucocytes falling to as low as 3,600 per c.mm. 
The leucopenia is most marked at the acme of the fever, and then 
diminishes gradually. There is also a decrease in coagulability of 
the blood. 

In general the patients complain of headache, dizziness, tinnitus, 
partial deafness, etc. In severe cases there is delirium, stupor or 
coma. In some cases stiffness of the neck occurs, and the muscles, 
especially those of the extremities, become very sensitive to pressure. 
The knee-jerk, skin and cremasteric reflexes are usually present. 
Nervous symptoms generally disappear after subsidence of the fever, 
while in grave cases they remain after the apyrexia, a peculiarity of 
the disease. 

Relapses have not been observed, although in some cases high 
fever returns after defervescence; such cases are usually fatal. 

As to reinfection, I have observed several cases, two with an 
intermission of a few months, and one after about two years. 
Recovery from a first attack generally ensures a relative immunity, 
as a second attack is milder, and natives and early immigrants are 
less liable to infection. 

Among seventeen deaths, six occurred in the second, eight m the 
third, two in the fourth and one in the seventh week. 

The death rate varies greatly according to locality and year. 
It amounted to from o per cent. (1914) to 3 per cent. (1915) in the 
Toyoda plantation, and from 15 per cent. (1914) to 12 per cent. 



(1915) to the Hayashida plantation. In the Karenko district the 
death rates were 10 5 per cent. (1914), 9*3 per cent. (1915)1 and 
ii*66 per cent. (1916). 


B. Treatment 

The site of the bite should be treated quickly and energetically, 
best by extirpation or cauterisation, otherwise the treatment is 
symptomatic only. 

C. Diagnosis and Differential Diagnosis 

A typical case of the disease may easily be recognised by the 
presence of an ulcer, buboes, and fever with exanthemata. 

From typhus fever, with which it has a certain similarity 
especially in the rash, it can be distinguished by the buboes. 
Incidentally, it is worth noting that in Formosa, typhus fever is 
unknown up to date. 

In dengue fever, which is sometimes epidemic in the island, 
glandular enlargement is exceptionally observed, but the ulcer is 
unknown. In measles, the nature of the rash, the absence of ulcer 
and bubo, as well as its epidemic character, enable a diagnosis to be 
made. 

From venereal ulcers and bubo, the present disease may be 
differentiated by the character of the ulcer, by the bubo and 
exanthem, and more especially by the history. 

From other ulcers, lymphangeitis and adenitis, caused by 
suppurative cocci, a diagnosis can be established bv microscopical 
examination as well as by the appearance of the rash. 

Weil’s disease, caused by Leptospira icterohaemorrhagiae , 
Inada, is always accompanied by icterus, and has never been 
observed in this island 

Seven-day' fever, or the autumnal fever of Fukuoka and 
Shizuoka-ken, is a spirochaetal disease caused by Leptospira 
hebdomadis , Ido, one suspected case of which was observed by 
Dr. Ohnishi at Taihoku. These are differentiated from the 
tsutsugamushi disease by the presence of icterus and the causal 
spirochaetae or by serum reactions. 



250 


IV, SOME REMARKS ON THE JAPANESE AND FORMOSAN 
TSUTSUGAMUSHI DISEASE AND ALLIED ENDEMIC 
GLANDULAR FEVERS 

Though the identity of the endemic fever of northern Japan and 
Formosa, as can be seen in my earlier papers, is recognised, there 
still exists some differences between them, which may be summarized 
as follows: — 

(1) The mortality of the tsutsugamushi disease in Niigata and 
Yamagata districts in 1917 is estimated at 30 per cent, and 
50 per cent, respectively, while that of the Formosan fever amounted 
in recent years to about 10 per cent, per annum. 

(2) The Japanese fever appears regularly in the hot season from 
July towards the end of September, the mean temperature in these 
months being above 7i'2°F. at Nagaoka in the Niigata district. 
The Formosan fever generally begins in April and disappears in 
November.* The free season in the Yoshino plantation is in the 
months with a mean temperature below 75*0° F. 

(3) The endemic areas of Japanese fever are strictly restricted 
to certain parts of river banks and alluvial deltas or islets formed 
in the river, which are subjected to annual inundation, as the name 
‘Japanese flood or river fever* given by Prof. E. Baelz indicates. 
In Karenko district, however, the endemic areas are situated in 
river valleys or fertile plains and woodlands, either at the foot of 
mountains or on elevations free from inundation. 

(4) The host of the tsutsugamushi or akamushi mite of Japan 
is the field mouse, Microtus montebeUi; those of the Formosan mite 
are common rodents and insectivora and other wild or domestic 
animals, e.g., chickens, pheasants, and dogs. 

(5) While the exanthem occurred in nearly all of the cases in the 
Yamagata district (Ohnuma and Naganobori), it is absent in 
5 per cent, of Niigata cases (Commissioner’s report), and thirty-five 
non-exanthematous among one hundred and four cases were 
recorded in 1915 in the Karenko district. 

(6) The respiratory system was affected in 48 per cent, of the 
cases in Niigata; in 34 per cent, of Formosan cases in 1915. 

(7) It is said that the Japanese monkey is susceptible to the virus 

• It if reported from Ako district office that four typical cases of the disease were observed 
hv a local physician in January, IQ19. 



251 

of the tsutsugamushi disease, but this is not the case with the 
Formosan virus. 

(8) The virus-transmitting red mite of Japan and Formosa is 
the larva of Trombicula akamushi, Br. 

Dr. Schiiifner has observed, since 1902, a special type of fever in 
Deli, Sumatra. He gave the name ‘ pseudo-typhus ’ to the disease, 
and believes it to be a variant of the tsutsugamushi disease. The 
following is a summary of his paper (1913): — 

‘ In Deli, pseudo-typhus, with a mortality of 3 per cent., is 
found throughout the year. So far as is known, people 
contract the disease through the bite of Trombidium larvae. 
In a quarter of an hour after the bite, violent itching is felt. 
An initial ulcer occurs in the groin, thigh or neck in most 
cases. The adjacent lymphatic glands swell to the size of a 
pigeon’s egg. A characteristic eruption appears on the second 
or third day, develops for several days, and then slowly 
disappears. It occurs in 70 per cent, of the native cases. 
The course of the fever resembles that of typhoid fever, but it 
begins and terminates more acutely. The blood shows on an 
average a leucocytosis of 12,000, in which the increase of the 
lymphocytes and decrease of the polymorphonuclears is 
remarkable. Inoculation experiments with monkeys gave no 
result.’ 

Ashburn and Craig noted a similar disease in the Philippine 
Islands in 1908, but gave no information about the mode of 
transmission. 

Noc and Gantron reported two cases of undetermined fever 
observed at the Saigon Military Hospital in the autumn of 1914. 
These cases were very similar to those seen by Schiiifner at Deli. 

Dowden reported a suspected case of Kedani river fever observed 
in the Federated Malay States, viz., that of an European, aged 
20 years, admitted into hospital on the fourteenth day of the disease 
in the typhoid state. A profuse and haemorrhagic rash appeared 
on the eighth day. Glandular enlargement was marked, and there 
were neuritic symptoms. 

Weir reports the disease and the red mite from Corea. 

The Mossman fever which occurs in the Mossman district, North 
Queensland, and affects cane-cutters, etc., was first reported by 



252 


Smithson in 1910, and recently studied by Drs. Breinl, Priestly and 
Fielding; it has a great similarity to tsutsugamushi disease and 
Schiiffner's pseudo-typhus. It is characterised by an irregular 
remittent fever from three days to three weeks, accompanied by 
painless swelling of lymphatic glands and the appearance of a 
macular or vesicular rash. 

The above-mentioned endemic fevers, the Sumatran pseudo¬ 
typhus, Australian Mossman fever, and Philippine, Saigon and 
Malay fever, are exanthematous glandular fever, having many 
features in common. They are most probably caused by the bite of 
mites. 

Two cases resembling tsutsugamushi fever, said to have been 
caused by the bite of a tick, Antblyomma sp., have been reported 
from Karenko. Whether or no the cases were truly tick fever or 
tsutsugamushi disease due to mite bite could not be determined. 


VI. PROPHYLAXIS 

As the mode of infection of the endemic disease of Formosa has 
been clearly defined, preventive measures based on its aetiology 
should be adopted. 

(1) All parts of the body should be, in a mite-infested area, 
properly protected. Prof. Hayashi's or Prof. Nagayo’s mite-prool 
suit is recommended for this purpose. 

(2) All articles of clothing, including loin-cloths, used in an 
infected field, should be taken off outside the dwelling-house, and 
be fumigated with sulphur or boiled or dipped into a disinfectant 
solution. 

(3) At the same time, hot baths with the use of soap should be 
taken regularly. Vleminkx’s solution, Kummerfield's mixture, 
1 disinfectol' solution, etc., may be applied to those parts of the body 
liable to mite bite. 

(4) The skin of the body should be carefully examined after field 
work has been done. Examine with a lens any red spot for a red 
mite in its centre. If the mite is found, the spot should be excised 
or the mite should be removed with a needle. 

(5) The best prophylactic measure is to bring the infested areas 



253 


under cultivation. Before doing so, it is indispensable to bum 
all bushes and grasses. The clearing of the areas must be done 
during the free season. 

(6) It is very important to educate the inhabitants of the infested 
localities as to the nature of the disease and its transmitter. 


REFERENCES 


Ajcagi, K, and Riv, T. (1918). On a cate of Ttuttugamuthi Disease occurring in Kagi district, 
Formosa. Taiwan Igakkai Zassbi , No. 182-3. 

Ashburn, P. M., and Craig, C. F. (1908). A comparative study of Tsutsugamushi Disease 
and Spotted or Tick Fever of Montana. Philippine Journ. of Sci. B. Vol. Ill, p. 1. 

Brxinl, Priestlxy, and Fiklding (1914). On the occurrence and pathology of endemic 
glandular fever, a specific fever occurring in the Motsman district of North Queensland. 
Med. Journ. of Australia. Vol. i, p. 391. 

Dowdkn, R. (1915). A suspected case of Kedani River Fever in the Federated Malay States. 
Indian Medical Gazette. Vol. L, p. 208. 

Hatori, J. (1915-18). Reports I-IV on the Exanthematous Bubonic Fever (Tsutsugamushi 
Disease), endemic in Formosa. Taiwan Igakkai Zassbi. Nos. 147, 150, 170, 182. 

- (1916). On the Exanthematous Bubonic Fever of Formosa, and its identity with the 

Tsutsugamushi Disease of Japan. Nippon Eisei Gakkai Zassbi. Vol. XI. 

Hayashi, N. (1917). Piroplasma-like bodies as the cause of the Tsutsugamushi Disease. Kyoto 
Igaku Zassbi. Vol. XIV. 

Hayasbi, N., and Mukoyama, T. (1918). Studies on the Tsutsugamushi Disease in 1917. 
Cbuo Igakkai Zassbi. No. 138. 

Hokuttsu Igakkai (1904). The First Report on the Investigation of the Tsutsugamushi Disease. 

Kato, S. (1910). On an Exanthematous Fever observed in Kagi district Taiwan Igakkai 
Zassbi. No. 100. 

Kawamura, R., Arima, E., and Hattori, T. (1913). On the Pathological Anatomy of the 
Tsutsugamushi Disease. Nippon Byori Gakkai , Trans. Vol. IV. 

- (1914). On the Blood Changes of the Tsutsugamushi Disease. Hokuetsu Igakkai 

Zassbi. 29th annual set. 

Kawamura, R., and Hattori, T. (19x5). Histological Studies of the Exanthem of Tsutsu¬ 
gamushi Disease. Hokuetsu Igakkai Zassbi. 30th Annual Set 

-(1916). On the Causative Virus of the Tsutsugamushi Disease. Hokuetsu Igakkai 

Zassbi. 30th annual Set 

Kawamura, R., and Komagata, K. (1916). On the Morphology of the Akamushi. Hokuetsu 
Igakkai Zassbi. 31st Annual Set. 

Kawamura, R., and Yamaguchi, M. (1916). On the Adult and Nymph of the Akamushi. 
Hokuetsu Igakkai Zassbi. 31st Annual Set 

- (1916). On the Anatomy of the Adult and the Life Cycle of the Akamushi. Hokuetsu 

Igakkai Zassbi. 31st Annual Set. 

- (1917). Researches on the Etiology of the Tsutsugamushi Disease. Hokuetsu Igakkai 

Zassbi. 32nd Annual Set 

Kitajima, T., and Miyajima, M. (1909-10). Studies on the Tsutsugamushi Disease. Saikingaku 
Zassbi. Report III-IV. 

-(*9*8). Studien ueber die Tsutsugamushi-Krankheit, I. Kitasato Arcb. Exper. Med. 

Vol. II. 



2 54 


Miyajima, M. (1917). On the Life Cycle of the Akamushi, Carrier of the Nippon River Fever. 
Kitasato Arcb. Exper. Med . Vol. I. 

Miyajima, M., and Okumura, T. (1917). Comparative Studies on the Japanese, Korean and 
Formosan Akamushi and Allied Species. Saikingaku Zassbi. No. 266. 

- (1916). Comparative Studies on the Larvae of Trombidiidae. Saikingaku Zassbi. 

No. 254. 

- (1918) On the Akamushi Body, or Clamydomycetes Tsutsugamushi. Saikingaku Zassbi 

No. 269. 

Nagayo, M., Miyakawa, Y., Mitamura, T., and Imamura, A. (1915). On the Tsutsugamushi 
Disease. Tokyo Igakkai Zassbi. Vol. XXIX. 

- ( I 9 I 7). On the Nymph and Prosopon of the Tsutsugamushi ( Leptotrombidium akamushi ), 

Carrier of the Tsutsugamushi Disease. Journ. Exper. Med. Vol. XXV, p. 255. 

- ( I 9 I 7 )* On the Culture of the Causative Virus of Tsutsugamushi Disease. Ijisbinbun , 

No. 979. 

Nagayo, M., Miyakawa, Y., Mitamura, T., and Tamiya, T. (1918). On the Causative Virus 
of Tsutsugamushi Disease in the body of the adult Tsutsugamushi Mite. Densenbyo 
Kenkyujo Gakuyukai Zassbi. Vol. II, No. 1. 

Nakagawa, K. (1913). On an Exanthematous Fever prevailing at Karenko District. Taiwan 
Igakkai Zassbi. No. 125. 

Noc, and Gantron (1915). Deux casde fievre indetermince rappclant lc pseudo-typhus de Deli 
observes a Saigon. Bull. Soc. Med. Cbir. Indo-Cbina. Vol. VI, p. 108. 

Ogata, M. (1911-17). II-VIII Mittheilungen ueber die aetiologie dcr Tsutsugamushi 
Krankhcit. Mittbeilung d. Med . Facult. d. Kais.-Jap. Univ. z. Tokio. Vol. X-XVII. 

- (1918). On the Tsutsugamushi-eumycetes. Eiseigaku Densenbyogaku Zassbi. Vol. XIII, 

No. 6. 

Ohnishi, S. (1918). On the Seven-days Fever observed in Formosa. Taiwan Igakkai Zassbi. 
No. 186. 

Sano, Y. (1914). On the undetermined Exanthematous Fever. Taiwan Igakkai Zassbi. 
No. 138. 

Schuetfner (1913)- Pseudotyphus in Deli, Sumatra. Comptes rendus des traveax du Troisihme 
Congrcs Biennal , Saigon, p. 309. 

Senouye, Y. (1917). On a case of Tsutsugamushi Disease infected in Mt. Ari region. Taiwan 
Igakkai Zassbi. 

Tanaka, K. (1899). Ueber die Aetiologie und Pathogencse der Kedani Krankhcit. Ccntralbl. 
f. Bakt. u. infect. Kr. I. Abt. Bd. XV 7 1 . 

- (1906). Ueber mcine Japanische Kedani Krankhcit. Ibid. Bd. XLII. 

- (1918). Differentiations of the Kedani, pseudo-akamushi, Japanese Leptus autumnalis 

and its adults. Ikaijibo. No. 1228. 

Tsuruki, T. (1915). The Tsutsugamushi Disease in Yamagita-ken. 

Yersin, A., and Vassal, J. J. (1908). Typhus fever in Indo-China. Philippine Journ. Sci.. 
B. Vol. Ill, p. 131. 




2$6 


EXPLANATION OF PLATES 

Plate VIII 

Fig. i. View of Mokkui Valley at the time of the expeditionary 
campaign, 1914. 

Fig. 2. Larva of Tr. akamushi. x 100. 

Fig. 3. Nymph of Tr. akamushi. 

Fig. 4. Adult of Tr. akamushi. 









258 


Plate IX 

Fig. 1. An ulcer on scrotum caused by bite of virulent akamushi. 

Fig. 2. Patient in acute stage [after Dr. Ohta], D , denotes 
ulcer; o, inguinal bubos. 








259 


NOTES ON THE BIONOMICS OF 
STEGOMTIA FASCIATA , Fabr. 

(Part I) 

BY 

J. W. FIELDING 
Australian Institute of Tropical Medicine 
{Received for publication 18 th August , 1919) 

INTRODUCTION 

Since the discovery of the role of Stegoniyia fasciata in the 
transmission of yellow fever, much attention has been devoted to the 
study of the life history and habits of this mosquito in Europe, 
Africa and America. 

Notwithstanding the vital importance of such knowledge to 
Australia, no research work of any importance, other than Taylor’s 
inquiry into the distribution of the species in the coastal towns of 
Queensland, has been undertaken in Australia, either to confirm or 
to supplement the result of investigations carried out elsewhere by 
Finlay, Goeldi, MacGregor, Carrol, Reed, Newstead, Boyce, Bacot 
and many other workers, to whom we are indebted almost entirely 
for our knowledge of the bionomics of this widely distributed species. 

The necessity for breeding large numbers of Stegomyia fasciata 
to meet the requirements of this Institute appeared a favourable 
opportunity to undertake some of this hitherto neglected work. 
Although very incomplete, it has been considered advisable to 
record the results of these observations and experiments at this stage, 
rather than to defer their publication until further contemplated 
experiments have been undertaken. 

It will be observed that where similar information has been 
sought, I have followed very closely the methods and technique of 
BaCot (1916), whose paper has been freely consulted, as being by far 
the most complete and comprehensive of its kind yet published. 



26o 


The various stages of Stegomyia fasciata have been so fully and 
accurately described by earlier writers that it is quite unnecessary to 
attempt to add anything to their descriptions, nor can new facts be 
added regarding the seasonal prevalence and distribution of this 
mosquito in Townsville to the published records of Taylor (1915). 

The accompanying chart shows the temperatures and humidity 
readings taken in the Institute grounds during the period in which 
the experiments were carried out. 



METHODS OF KEEPING AND FEEDING ADULT MOSQUITOES 

In these experiments the adult mosquitoes were kept in small 
cages, as commonly used, measuring 12 by 8 by 7 inches, and 
consisting of a light wooden frame covered with cheese cloth on three 
sides and one end, and zinc on the bottom. The remaining end was 
fitted with a calico sleeve sufficiently large to permit of the passage 
of the arm and necessary water containers. A glass panel occupied 
one-third of one side of each cage, but it was considered to be of no 





26 i 


advantage when light fabric was used for screening the remainder 
of the cage. The mosquitoes were offered food daily, the following 
substances being tried at different times, viz., blood, banana, 
peptone and sugar solution, milk and sugar, etc. After some initial 
trials blood was found to be the most satisfactory for the females and 
banana for the males, these foods being used subsequently to the 
exclusion of all others. 

Feeding experiments with small animals, such as guinea-pigs, 
rats and puppies, were, as a rule unsuccessful, even when the animal 
was left in the mosquito cage overnight. The most satisfactory 
method of feeding* with blood, and the one adopted throughout the 
following experiments, was found to be the presentation of the arm 
and hand each morning for a period of from ten to twenty minutes 
according to circumstances, it being noticed, for example, that 
during hot and humid weather the mosquitoes fed more readily than 
in cool weather. 

In some current experiments, in which it is desired to feed 
Stegomyia on dog's blood, successful results are obtained by the 
following method :—One of the cages previously referred to is placed 
upon the floor and the dog's head thrust into it, the animal being 
kept in a recumbent position and the sleeve of the cage secured 
behind its head with one hand whilst the other presses the body to 
the floor to prevent the animal rising. To ensure successful feeding 
it is found necessary to shave or clip the hair on the face and ears. 
Other methods of feeding upon dogs were tried, but found to be 
unsatisfactory. 


i. EGGS 

The maximum number of eggs laid by a fertilised captive 

female 

From a long series of observations it has been found that the 
average egg production of a fertilised captive female fed on blood 
is about seven hundred and fifty, covering a laying period of from 
forty to seventy-two days. None of my records cover a longer 
period of productiveness, although females have been kept in 
captivity up to a maximum of ninety-three days. It was found 
impossible, however, owing to absence from Townsville, to continue 



262 


blood feeding after the sixty-sixth day, banana being substituted for 
blood, with the invariable result that egg production ceased on the 
change of food. 

The following record may be cited as typical of many illustrating 
the rate of egg production. A female, bred for five generations in 
the laboratory, was segregated with two males and fed exclusively 
on human blood. The first feed was taken four days after her 
emergence, and the first batch of eggs laid three days later. During 
the first thirty-one days of her life she fed eight times and laid 
four hundred and thirty-seven eggs. During the succeeding 
thirty-one days she fed six times and laid two hundred and sixty 
eggs. Two more feeds were taken and fifty-five eggs laid during 
the following ten days. In all she fed sixteen times and laid seven 
hundred and fifty-two eggs during the seventy-two days of her life. 
On the day following the laying of the last batch of eggs she was 
accidentally destroyed. A complete record of these eggs was not 
kept to determine the percentage of fertility, but a certain proportion 
of the batches were kept under observation and found to produce 
between 80 per cent, and 90 per cent, of larvae. 

The selection of situation for oviposition 
(a) Females in captivity 

The receptacles usually provided in the cages for oviposition 
were dishes 3 inches to 4 inches in diameter, and contained about 
J-inch to 1 inch of water. The favourite position of the ovipositing 
female appears to be resting on the sides of the receptacle with the 
apex of the abdomen just touching the water. Very often eggs are 
laid on the surface film of the water as the female moves about on it, 
but occasionally as many as fifteen eggs are laid in a batch. 
A considerable proportion of the eggs laid near the margin of the 
re ceptacles are drawn against the sides by capillary attraction, and 
become stranded as the water dries. Under more natural conditions 
these doubtless form the bulk of the resistant eggs which lie dormant 
until the supply of water is replenished. When wet filter paper is 
provided the eggs are usually scattered over its surface, but 
occasionally they are laid so close together as to overlap. 



263 


( b ) Wild females 

It has been frequently noticed that almost any vessel containing 
water and left in or near the Institute building would attract the 
females of Stegomyia and offer a suitable place for oviposition. 

In view of the opinion held by some other observers that these 
mosquitoes show a decided preference for certain positions and 
certain rooms, the investigations detailed in Experiment No. 1 were 
undertaken with the object of confirming, or otherwise, the results 
of my casual observations. 


Experiment I 


No. 

Position 

Nature of Receptacle 

Contents 

Stegomyia 
eggs laid 

Remarks 

1 

Workroom on floor 

Enamel bucket 

Tap water . 

+ 


2 

Workroom on bench 

Earthenware sink 

Tap water ... 

-r 


3 

Workroom on bench 

Pickle bottle . 

Tap water and sea water 
equal parts 

4- 


4 

Workroom on bench 

Pickle bottle . 

Rain water . 



5 

Workroom on bench 

Tin used as water bath 

Rain water . 

a- 

Also Culex sp. 

6 

Hallway . 

Glvd. iron cistern 

12 ft. from floor 

Rain water ... 

+ 


7 

Bathroom on bench 

Glvd. tin . 

Tap water . 

+ 


8 

Bathroom on floor 

Earthenware jar 

Tap water ... 

+ 

Also Culexfatigans. 

9 

Water Closet . 

Cast iron cistern not 
used for some time 

Tap water . j 

+ 


10 

Sterilising room on bench 

Enamel bowl . 

• Tap water and dirty tubes | 

+ 


11 

Ster. room under bench 

Earthenware jar 

Tap water and dirty slides 

+ ! 

Also Culex fatigans 
and C. ? sitiens 

12 

Animal house on shelf 

8 ft. from floor 

Glvd. iron dish 

Tap water . 

+ 


*3 

Animal house on shelf 

8 ft. from floor 

Enamel bowl . 

Tap water . 

+ 

I 

*4 

Under roof of Institute ... 

Glvd. iron dish 

Tap water. 

— 

1 

l 

*5 

Under roof of Institute ... 

Glvd. iron dish 

Tap water. 


! 

i 

16 

Under roof of Institute ... 

Enamel bowl ... 

Tap water ... 

- 

i_. 


The investigations on the whole tend to show that in Townsville, 
at any rate, Stegomyia do not appear to show a preference for 
certain rooms or situations, but will oviposit in almost any receptacle 
containing water. This finding is noteworthy in view of the fact 







264 


that Bacot’s experiments gave entirely different results. In a set of 
twelve receptacles under observation he.found Stegomyia ovipositing 
in only one—a card cream jar in the kitchen. It must be noticed, 
however, that he found Stegomyia ovipositing in such positions as 
the following: in safe stands in gallery, in a tin bowl in yard and 
in tins and wooden tub in the mosquito house. 


Baits to attract pregnant females 

Experiments to ascertain if ovipositing females prefer 
contaminated water to clear 

The following experiments (see Experiment II) were carried out 
in a cage containing a large number of Stegomyia , the females of 
which were given opportunities of feeding regularly on human 
blood. The glass dishes each contained 200 c.c. tap water, in 
addition to the organic matter, and were placed as far apart as the 
size of the cage would permit. Except in the case of leaves, which 
stood for forty-eight hours, none of the cultures were permitted to 
stand for more than twenty-four hours before being introduced into 
the cage. One trial was carried on a second day owing to the small 
number of eggs laid on the first. 

Bacot, who carried out similar experiments, remarks : * While the 
evidence of selection . . . appears definite, it is very far from being 
the unanimous unvarying character that might be expected from a 
deep-seated instinct coupled with larval needs. On the contrary, it 
carries with it a certain suggestion of bias on the part of the females 
in reference to their own tastes largely parallel, but not necessarily 
identical, with provision for their offspring. ... * 

It appears to me that these remarks might be fittingly applied to 
the results of mv experiments. It might be observed, however, that 
sugar and water appeared to be the most attractive bait, yet it has 
been shown that sugar gave unfavourable results as a larvae food. 



265 

Experiment II 




No. of 

Percentage 


No. 

Contents of dish 

eggs laid 

of total 

Remarks 


(1st Series) 




1 

Tap water . 

30 

8-6 

No smell 

2 

Tap water plus 0*5 grm. dried mango 




leaf, standing for 2 days . 

30 

8-6 

Did not smell strongly 

3 

Tap water plus 0*5 grm. fresh fowl 

120 

34-3 

Did not smell strongly 


faeces . 



4 

Tap water plus 0*5 grm. freshly killed 





cockroach . 

170 

48-5 

Smelled strongly 


Total eggs laid. 

35 ° 




(2nd Series) 




1 

Tap water . 

40 

6-66 


2 

Tap water plus 0*5 grm. dry pawpaw 





leaf allowed to decompose for 2 days 

20 

3*34 

Did not smell strongly 

3 

Tap water plus 0*5 grm. dried cockroach 

186 

V *oo 

Slight smell 

4 

Tap water plus icc. ox bile . 

160 

26*66 

Fairly strong smell 

5 

Tap water plus 0*5 grm. Sodium 



Fairly strong smell 


Taurocholate. 

194 

32*34 


Total eggs laid. 

600 



1 

(3rd Series) 





1st Count 




1 

Tap water . 

M 

21*5 

On account of the few 

2 

Tap water plus 0-5 horse faeces 

20 

30-8 

eggs laid the dishes 

3 

Tap water plus 1 cc. urine 3 days old ... 

3 1 

47*7 

were placed in the 
cage again on the 


Total eggs hid. 

6 5 


following day and a 
second count was 
made 48 hours later. 



(3rd Series) 

2nd Count 




1 

Tap water . 

120 

24-0 

Did not smell 

2 

Tap water plus 0*5 horse faeces 

90 

18*o 

Smelling 

3 

Tap water plus 1 cc. urine 3 days old ... 

290 

58*° 

Smelling strongly 


Total eggs laid. 

500 



1 

(4th Series) 




1 

Rain water . 

43 

20*5 

At the conclusion all 

2 

Tap water plus 0*5 grm. powdered rice 

40 

19*0 

dishes had a very 

3 

Tap water plus 10 grm. sugar. 

>27 

605 

slight smell 


Total eggs laid. 

210 




Table I contains a summary of this experiment (Experiment II). 





2 66 


Tabus i. 


Contents of Receptacles 

Percentage of the 
total eggs laid 
in each series 

Tap water and pawpaw leaf., 

3-34 

Tap water (2nd series). 1 

6-66 

Tap water (1st series). 

8-6 

Tap water and mango leaf . 

8-6 

Tap water and horse faeces (2nd count) . 

18*0 

Tap water and powdered rice . 

19*0 

Rain water . 

20*5 

Tap water (1st count). 

21*5 

Tap water (2nd count) . 

24-0 

'l ap water and ox bile. . 

26*66 

Tap water and horse faeces (1st count) . 

30*8 

Tap water and dry cockroach. 

31*0 

Tap water and sodium taurocholate. 

3 2 ‘34 

Tap water and fowl faeces . 

34*3 

Tap water and urine (ist count) . 

477 

Tap water and fresh cockroach . 

48-5 

Tap water and urine (2nd count) . 

580 

Tap water and sugar. . 

60*5 


Experiments to determine whether Stegomyia would 
oviposit on sea water 

For this purpose two buckets, each containing 4 litres of water, 
were exposed in a room where Stegomyia were fairly numerous. 
Bucket 4 A * contained sea water drawn from the harbour, and 
bucket 4 B * tap water from the town service pipes. The sea water 
was exposed continuously for a period of two months (19th April to 
19th June), during which time no eggs were laid on it, whilst the 
other containing tap water was exposed for one month (19th May to 
19th June) and was frequently visited by both Stegomyia and Culex 























267 

fatigans for oviposition. In all, four hundred and ninety-one eggs 
of Stegomyia were laid. 

A further series of experiments were then carried out to determine 
the amount of dilution necessary before Stegomyia could be induced 
to oviposit upon sea water mixed with rain water. For this purpose 
jars containing various percentages of sea water, from 10 per cent, to 
undiluted sea water, were exposed in one of the stock-breeding cages 
containing numerous females, which fed regularly on human blood. 
As each jar was found to contain eggs it was removed from the cage, 
and that portion of the experiment was considered finished. In the 
case of the jars containing 80 per cent, to 100 per cent, sea water, the 
vessels were kept under observation for a considerable period and 
until there was no further possibility of oviposition taking place. 
The results are shown in Table 2. 


Table 2. 

Number of eggs of Stegomyia fasciata laid on different percentages of sea water. 


Percentage of sea water 
in rain water 

1 

j Number of eggs laid 

Remarks 

10 


Numerous eggs were laid but 
were not counted 

20 

?J 


3 ° 

12 


40 

43 


5 ° 

2 4 


60 

28 


7° 

20 


80 

nil 


90 

nil 


100 

nil 



Hatching of Eggs of Stegomyia 

The period elapsing between the deposition of the eggs and the 
emergence of the young larvae is extremely variable. In the routine 
of breeding Stegomyia in this Institute, it has been found that the 



268 


great majority of eggs laid overnight in the small dishes of water 
provided for the purpose hatched during the second and third day 
following in summer, and during the third and fourth day in winter. 
A certain proportion of eggs of some batches, however, do not hatch 
within this period, but do so after a further period of immersion, this 
period being, as I have ascertained, so long as four and a half 
months. Had further observations been made in this direction, it is 
very probable that the period of delayed hatching of continuously 
immersed eggs would have been found to be sometimes longer, since 
Bacot has shown that under these conditions hatching may be 
delayed from two to five months. 


The influence of periods of drying upon the hatching 

of eggs 

Theobald (1903) first drew attention to the fact that eggs of 
Stegomyia fasciata are capable of withstanding the effects of long 
periods of dessication. Since then, several others have published the 
results of experiments to ascertain the maximum period of viability 
of these eggs under various conditions of dry storage, and to 
elucidate the problems concerning the factors which bring about their 
subsequent hatching. Bacot believes that the incubation period of 
the eggs in Freetown is from thirty to forty hours, but that hatching 
of any given batch may be distributed over a lengthy period if 
subjected to periods of drying. The whole of the eggs of a batch 
may hatch as soon as they are replaced in water, or a proportion of 
them only may do so. Others may resist the first or second 
immersion and yield to a subsequent one. The principal factors 
which act as stimuli to hatching appear to be temperature and 
humidity. As he points out, a response to cooling would well serve 
the needs of the species in allowing of the fullest advantage being 
taken of the facilities for breeding in small temporary accumulations 
of rain water. 

With the object of confirming the above observations under local 
conditions a number of experiments (III, IV, V) were carried out, 
which will be found to accord generally with the conclusions arrived 
at by earlier investigators. 



269 


Experiments to show the effects on eggs of drying and 
replacement of water 

Experiment III 


Twenty eggs laid overnight were placed in a tube inches in 
diameter, containing 15 c.c. of tap water, which completely 
evaporated and was replaced on the days shown. 


1 

Days . 

3 

5 

7 

8 

11 

14 

*5 

20 

21 

Total 

Number of eggs hatched. 

2 

4 


9 

2 


1 



18 

Day* when found dry (indicated — ) 

... 

- 

- 



- 


- 


... 

Days when water replaced (indicated -f) 




4- ; 



4- 


4- 

... 


Thus eighteen eggs (90 per cent.) hatched in fifteen days and 
after two dryings; two eggs (10 per cent.) were found to be collapsed 
when examined on 24th day. 


Experiment IV 


Twenty-five eggs laid overnight were placed in a petri dish 
3 inches in diameter, containing 10 c.c. of tap water, which 
completely evaporated and was replaced on the days shown. 


Day* . 

| 

3 

4 

5 

6 

8 

i: 

II 12 

1 5 

18 

Number of eggs hatched. 

3 

8 



5 

i 


... i ... 



Days when found dry (indicated —) 



- 

... 


- 

- 

... , - 

- 

- 

Days when water replaced (indicated 4 ) 




4- 


4- 

4- 

... 4- 

+ 

4- 


Experiment IV ( continued ). 


Days . 

20 

21 

, 1 

23 26 j 

1 

1 

28 

1 ! 

29 1 

32 

i 

33 

34 

Total 

Number of eggs hatched. 

i 

7 

1 J 

. 




1 

25 

Day* when found dry (indicated — ) ... 

- 

... j 

- 1 ... 

- 


- 



... 

Days when water replaced (indicated 4-) 

4- 


4* ... 

l 1 


4- 


4- 




Thus 100 per cent, of the eggs hatched in thirty-four days and 
after ten dryings and ten replacements of water. 





27 ° 

Experiment V 


Twenty-eight eggs laid overnight were placed in a petri dish 
4 inches in diameter, containing 15 c.c. of tap water, which 
completely evaporated and was replaced on the days shown. 


Days . 2 

3 

l 

l 

4 

5 

7 

9 

ii 

*3 

16 

*7 

20 

22 

Number of eggs hatched . 6 

... | 

l 3 




... 

... 

... 

... 

... 

... 

Days when found dry (indicated — ) 

i 


- 

- 

- 

- 

- 

- 


- 

- 

Days when water replaced (indicated + )j .»* 

+ 


-f 


+ 

i 

, + 

+ 


+ 

+ 

... 


Experiment V ( continurd ) 


Days . j 23 

! , 
25 j 26 

27 

20 

31 

32 34 


Total 

Number of eggs hatched.. 

Days when found dry (indicated — ) ... ... 

Days when water replaced (indicated -f) + 

| 1 

... | + 



+ 

j 

... | + 

4 

+ 

25 


Thus twenty-five eggs (89‘2 per cent.) hatched in thirty-eight 
days and after thirteen dryings and replacements of water. The 
remaining three eggs were found to be uncapped on the thirty-ninth 
day, but no larvae were seen. Assuming that these three eggs were 
uncapped by larvae, 100 per cent, of the eggs in this experiment may 
be said to have hatched. 

The viability of eggs after long periods of dry storage 

The longest period during which the eggs of Stegomyia fasciata 
are known to have remained viable in dry storage appears to be 
two hundred and sixty-two days (Bacot (1916)). For his experi¬ 
ment, Bacot collected eggs laid in Sierra Leone in January, from 
whence they were sent to England and immersed on 20th October. 
Within two hours of immersion the eggs hatched freely. 

The possibility of the change of climate and environment 
influencing the period of viability, as shown by Bacot’s experiment, 
suggested the desirability of carrying out some experiments wholly 
in Townsville. In one test (Experiment VI) eggs were kept in a 
viable condition in dry storage for a period of two hundred and 
fifty-seven days. In two other tests eggs retained their viability for 



271 


over two hundred days (216 and 218), and in five other tests for 
over one hundred and fifty days (198, 172, 169, 163, 154). The eggs 
used in Experiment VI were taken from the stock-breeding cages 
and were known, from other experiments, to be of normal fertility. 
Those in * A * were stored on a shelf in the laboratory, excepting in 
the case of ‘ A * 7, which was placed between the iron roof and the 
ceiling of an upstairs room of the Institute building. The eggs used 
in ‘ B ’ were stored on a layer of sand on the roof of an out-building, 
where they were protected from the rain by a flat sheet of iron 
resting on four legs about 18 inches high. Since Bacot and others 
have shown that in order to withstand long periods of drying eggs 
must pass through a preliminary incubation period in contact with 
water, all the eggs used in ‘ A 1 and 4 B * were incubated as shown in 
Experiment VI. 

Experiment VI 


A. Indoor?. 


No. 

No. of 
eggs laid 

Date of 
laying 

Maximum 
possible time 
in contact 
with water 
before storage 

Nature of storage 
receptacles 

Date of 
immersion 

No. of eggs 
hatched 
after 1 
immersion 

Viable 
for days 

Percentage 
of eggs 
hatched 

1 

24 

30.4.17 

Hours. 

2 5 

Muslin covered petri dish 

28.8.17 

14 

120 

5 8 ’3 

2 

. 25 

I * 5* , 7 

2 7 

Muslin covered petri dish 

28.8.17 

2 

”9 

8 -o 

3 

154 

5 - 5-17 

33 

Muslin covered tube 

28.8.17 

1 

32 

1 *4 

20-8 

4 

68 

8 - 5-*7 

35 

Glass covered petri dish 

28.10.17 

! 33 

* 7 2 

48-5 

5 

9 2 

, 3 - 5- I 7 

40 

Glass covered petri dish 

28.10.17 

40 

i6 7 

435 

6 

61 

2 3 - 5- , 7 

9 1 

Glass covered petri dish 

28.12.17 

3 2 

216 

5 2 ’4 

7 

200 

(approx.) 

29.4.17 

1 *3 

Glass covered petri dish 

11.1.18 

2 

2 57 

i-o 

(approx.) 


B. Outdoors. 


1 

4 * 

7 - 4-17 

43 

Muslin covered tube 

28.8.17 

2 5 

' 

1 42 

60-9 

2 

108 

4 - 5-17 

4 1 

Muslin covered tube 

28.8.17 

2 5 

1 *5 

2 V l S 

3 

7 ° 

I 7 ' 5* , 7 

40 

Gelatine capsule. 

28.10.17 

2 9 

163 

41-4 

4 

44 

2 3 * 5* I 7 

!I 3 

Muslin covered tube 

28.10.17 

2 

1 54 

4*5 

5 

62 

2.6.17 

138 

Glass covered dish 

11.1.18 

1 

218 

i-6 

6 

110 

23.6.17 

109 

Muslin covered dish 

11.1.18 

1 

198 

0-9 






272 


In a further series of experiments to determine the maximum 
period of viability after long periods of drying, thirty-two batches of 
eggs, numbering from thirty to one thousand, were stored for 
periods varying from three hundred and ninety-one days to 
five hundred and seventy-nine days and then subjected to stimuli to 
induce hatching. In all of these experiments negative results were 
obtained. 


Resistance of eggs to drying over Calcium chloride 

A number of experiments were carried out to determine the effect 
of Calcium chloride on eggs. Apparently this method of drying was 
too severe. The results are shown below: — 

Eggs dried for 7 days ... 80 to 90 per cent, of the eggs hatched. 

,, ,, ,, 19 n ••• ^ to 15 per cent. ,, ,, ,, 

,, ,, ,,26 ,, ... None hatched. 


Stimuli to the hatching of resistant eggs 

Cooling of resistant eggs as stimulus 
to hatching 

Bacot has shown that a rapid fall of a few degrees in temperature 
acts as a decided stimulus to the hatching of eggs which have been 
dried and have resisted subsequent immersion, or which have resisted 
the influence of the addition of fresh water. 

In Experiment VII will be found the results of investigations 
which appear to fully confirm the first of these findings, although in 
these the eggs were not first subjected to periods of wetting and 
drying. In this experiment eggs were collected from the stock- 
breeding cages and kept in water until placed in the ice chest. Those 
eggs which failed to hatch within three days were considered to be 
'overdue,* this ‘overdue* period being shown below. Just before 
cooling, the eggs were placed in a dish containing 200 c.c. of tap 
water and removed to the ice chest, where they remained from two 
to forty-eight hours as shown, the temperature falling from about 
8o°F. to between 51*8° and 57*2° F. 



2 73 


Experiment VII 


1 

Expt. 

1 

Eggs laid i 

i 

1 

Cooled 

Days 

Overdue 

No. of eggs 

No. hatched 
within 

2 hours 

No. hatched 
within 

24 hours 

No. hatched 
within 

48 hours 

Percentage 

hatched 

i 

25.3.17 

3 - 4- ! 7 

6 

6 

6 


1 

! 

1 oo-o 

2 

I 3 - 4 -I 7 

24.4.17 

8 

16 

10 



62-5 

3 1 

17.4.17 

2.5.17 

12 

6 

2 



33-3 

4 

4 - 5- ! 7 

26.5.17 

*9 

35 

8 

5 

*3 

76-0 

5 

8.6.17 

27.6.17 

16 

24 

*4 



5»*3 

6 

13.6.17 

27.6.17 

11 

8 

6 


1 

75-0 

7 

19.6.17 

27.6.17 

5 

108 

45 


! 

4*7 

8 

2 .1.l8 

18.1.18 

13 

25 

H 

4 

i' ' 

76-0 


Thus in this experiment of two hundred and twenty-eight eggs 
which failed to hatch during continuous immersion in water for 
periods varying from five to nineteen days longer than is generally 
required by non-resistant eggs, one hundred and twenty-eight were 
induced to hatch after periods of cooling ranging from two to forty- 
eight hours. Of these one hundred and twenty-eight eggs, one 
hundred and five hatched within two hours of being placed in the ice 
chest, nine within twenty-four hours, and fourteen within forty-eight 
hours. 


The effect of lysol on resistant eggs 

In certain experiments carried out at this Institute, it was found 
necessary to sterilize the eggs of Stegomyia , lysol being selected as a 
suitable agent. It was noted that after a short period of immersion 
in the disinfectant, resistant eggs hatched out soon after being 
submerged in water. This finding suggested the following experi¬ 
ment (VIII). 

The eggs used were from the stock-breeding cages, and proved 
to be of average fertility. They remained in or on water from the 
time of laying until immersed for thirty seconds in lysol (of the 
strength retailed by the manufacturers), the overdue period being 
reckoned from the third day after laying to the date of immersion 



274 


in the disinfectant. After treatment in lysol, the eggs were drained 
on filter paper and then immersed in water, the controls remaining 
in water throughout. The results of this treatment are shown in 
Experiment VIII. 


Experiment VIII 


Expt. 

No. of eggs 
treated 

No. of 
control eggs 

Date of 
treatment 

No. of 
days over¬ 
due in 
hatching 

No. of eggs hatched 
inside 2 hours. 

No. of eg 
in 24 

gs hatched 
hours 

Percentage 
hatched 
after lysol 

1 

13 

3 

20.5.17 

11 

Treated 

7 

| Control 

0 

Treated 

5 

Control 

0 

93*3 

2 

2 

3 

28.5.17 

7 

2 

0 

... 

0 

1 oo-o 

3 . 

2 

3 

N 

OO 

V| 

7 

2 

0 


0 

100*0 

4 

5 

3 

28.5.17 

7 

2 

0 

2 

0 

8o*o 

5 

3 


28.5.17 

6 

2 


1 


100*0 

6 

3 


29.5.I7 

8 

2 

... 

1 


100-0 

7 

i 

7 

: ' 

I 

4.6.17 

7 

7 

0 1 


1 

100*0 

8 

5 

7 

5.6.I7 

8 

3 

o ; 

2 

0 

100*0 

9 

1 

5 

7 . 6.17 

10 

2 


... 

1 

100*0 

10 

4 i 


8.6.I7 

11 

2 

° 1 

2 


100-0 

11 

6 

6 

11.6.17 j 

14 

j 

0 1 


0 

50-0 

12 

3 

3 

14.6.17 ! 

1 

>7 

3 

0 l 


0 

100*0 

l 3 

3 


15.6.17 ; 

[ 

18 i 

1 




33*3 


In this experiment (VIII), fifty-eight resistant eggs were treated 
with lysol, whilst thirty-four were used as controls. Of the former, 
thirty-eight hatched within one to two hours following their 
re-immersion in water after the lysol treatment, and thirteen others 
followed during the succeeding twenty-four hours. In all 879 
per cent, hatched. Of the thirty-four controls, none hatched during 
the first one to two hours following the commencement of the 
experiment, and two only during the first twenty-four hours. 

The effect of 2 ’5 per cent . soap solution on resistant eggs 

Bacot, experimenting with larvicides, found that petroleum 
soft soap emulsion 1-8000, acted as a stimulus to the hatching of 




275 


resistant eggs in a similar manner to lysol. The present observa¬ 
tions show that soap solutions without petroleum also have a similar 
action. 

In my experiment (Experiment IX) eggs were taken from the 
stock-breeding cages and kept continuously in or floating upon 
water up to the time of their immersion in the soap solution. The 
overdue period was reckoned, as in the former experiments, from 
thiee days after laying to the time of immersion in soap solution. 
The eggs were placed in the solution without preliminary drying 
and, after about five minutes' immersion, were returned to tap water. 


Experiment IX 


Expt. 

No. of egg* 
treated 

No. of 
control eggs 

j Date of 
treatment 

No. of 
days over¬ 
due in 
hatching 

No. of egg* hatched 
inside 2 hours 

No. of egg* hatched 
in 24 houn 

Percentage 

1 

4 

3 

20.6.17 

*3 

Treated 

3 

Control 

0 

Treated 

1 

Control 

0 

lOC’O 

2 

4 

3 

20.6.17 

*3 

1 


0 

0 

25-0 

3 

5 

2 

20.6.17 


1 

0 

0 

0 

20-0 

4 

6 

1 


22.6.17 

17 

5 


0 


* 3-3 

5 

4 

3 

22.6.17 

20 

3 

0 

0 

0 

75 -o 

6 

! 

6 

3 

24.6.17 

20 

4 

0 

i 

0 

0 

667 

7 

5 

5 

25.6.17 

23 

4 

0 

0 

1 

8 o-o 

S 

4 

5 

25.6.17 

•*3 

4 

0 


0 

1 oo-o 

9 

5 

5 

25.6.17 

23 

3 1 

0 

0 

0 

6o*o 

10 

10 


27.6.17 

25-27 

7 


0 


70-0 


Thus fifty-three resistant eggs were treated with soap solution 
and twenty-nine used as controls; thirty-five of the former hatched 
within one to two hours of their re-immersion in tap water following 
five minutes' immersion in the solution, and one hatched during the 
twenty-four hours following the treatment. In all 69*2 per cent, 
hatched within twenty-four hours. Of the seventeen which failed 
to hatch, two (one in No. 2 and one in No. 3) were found to be 
infertile and fifteen to contain either living or dead larvae. Only one 
of the control eggs hatched. 



The influence of submergence of eggs upon hatching 
Experiment X 

In order to ascertain the difference, if any, in the hatching 
period of submerged eggs and those floating on the surface film, 
forty-five eggs laid during the night of 28th April were divided 
into two batches on the following morning, ‘ A * containing 
twenty-two eggs and ‘ B * twenty-three eggs. The former (‘ A *) 
were submerged on filter paper in a dish containing 300 c.c. of 
tap water, whilst the latter (‘ B *) were allowed to float on the surface 
film of a similarly prepared dish. 

Batch 1 A * (submerged) hatched as follows: eighteen on the 
third day and one on the twenty-third day. The three remaining 
eggs were taken from the water, dried and dissected on the twenty- 
eighth day, when they were found to be collapsed. In batch ‘B* 
(floating), twenty-one eggs hatched on the third day and one on 
each of the fourth and fifth day. 

From this one experiment it would appear that the position of 
the eggs in the containers, i.e., whether floating or submerged, has 
little influence upon the hatching, since in ‘A* 8r8 per cent, of the 
total eggs, i.e., 947 per cent, of the total fertile eggs in the batch, 
hatched on the third day, and in ‘B’ 91*3 per cent, of the total. 
This view is supported by observations during the routine work of 
breeding Stegomyia. 


The effect of sea ivater upon the hatching of eggs 
On dry storage eggs 
Experiment XI 

Three hundred eggs, after having been dry stored for a period 
of three weeks, were divided into two equal batches, * A* being 
placed in 300 c.c. of sea water and ‘B* in 300 c.c. of tap water. 
The effects of this immersion are shown in Table 3. 



. .*• 


277 


Table 3. 


“ A” 

In tea water 

“ g M 

In tap water 

Total number of eggs hatched at the end of: — 

Total number of eggs hatched at the end of:— 

15 minutes.. 

20 

15 minutes . 

nil 

30 minutes. 

40 

30 minutes . 

3 ° 

60 minutes. 

96 

90 minutes . 

34 

27 hours . 

no 

2 hours. 

70 



5 hours. 

*s 



27 hours. 

,2 S 

Percentage hatched 

••• 77*3 

Percentage hatched . 

833 


On freshly laid eggs 
Experiment XII 

Two hundred eggs which were laid during the previous night 
were transferred in the morning from the dish of fresh water to a jar 
containing sea water. After fifteen days’ immersion in sea water 
they were divided into ten batches of twenty eggs each, and placed 
in bottles, (1) containing tap water, (2) containing 10 per cent, 
sea water, (3) containing 20 per cent, sea water, and so on up to 
90 per cent, sea water. The subsequent history of these eggs is 
shown in Table 4. 

Table 4. 


No. 

Contents of bottle 

No. of eggs 
hatched 

Length of larval life or subsequent 
stage of development reached 

I 

Tap water 

*7 

16 adults reared 

2 

Tap water plus io%sea water ... 

5 

£ adults reared 

3 

V 20 °o ?? 

4 

4 adults reared 

4 

>1 3 °% ” 

5 

4 adults and 1 pupa reared 

5 

I 4 °°o ?? ••• 

2 

1 adult reared, 1 larva lived three day9 

6 

, r .Of 

I ** . 0 V 

> 

5 larvae lived three days 

7 

6o° tJ 

1 

lived one day 

8 

70 °'' 

, v* m 7 u /o n 

1 

lived under 16 hours 

9 1 

1 v » 80% ,, ... 

1 

lived under 16 hours 

10 

9 °°o ?? 

0 

— 












27 $ 


The effects on hatching of submergence in small or large 
quantities of water 

Bacot’s experiments to elucidate the above question failed to 
bring forth any very conclusive evidence, but it appeared from them 
that slightly increased mortality and delayed hatching followed as 
a result of submergence in a small quantity of water. Nor do the 
present experiments afford convincing evidence, but such as has 
been adduced is in support of Bacot’s finding regarding the 
increased mortality amongst eggs submerged in small quantities of 
water. 

In the first of these experiments, thirty eggs laid during the 
night of 25th May were divided into two batches and submerged on 
filter paper, ‘A’ in 300 c.c. and ‘ B’ in 1,300 c.c. of tap water. 

In the second, sixty eggs laid on 30th May were similarly 
treated. The results are readily seen in Tables 5 and 6. 


Table 5. 


Batch 

Day 

Mortality 

3 

4 

5 

6 

7 

8 9 

IO 

I I 

12 

1 

>3 

i 

1 '+ 

*5 

16 

*7 

18 

28 

‘A’ 

‘ B’ 

6 

8 

1 

2 

i 

I"’ 


t 

... j ... 

i 

2 

4 

i 

I 

i 

... 



... 

... 

... 

> 3 - 3 % 

nil 


The two remaining eggs in 1 A 9 failed to hatch by the twenty- 
eighth day. 


Table 6 . 


Batch 

Day 

Mortality 

3 j 4 

5 

6 

7 

8 

9 

10 

1 * 

12 

13 

! 4 

! 

15 ! 

1 

16 

1 i 

17 18 

*9 

20 

21 

22 

‘A’ 

!... . 25 



-> 

... 

... 


* 






1 

...,... 1 

... 

... 

... 

... 

6-6% 

‘ B’ 

i -! 20 


3 

4 



2 


1 




... | 

| 


... 

... 

... 

nil 


The two remaining eggs in ‘ A ’ which failed to hatch by the 
sixtieth day were dried and found to be collapsed (dead). 






2 79 

The influence of temperature and environment on the hatching 
of eggs and the development of the resulting larvae 

In Experiment XIII, ninety eggs recently laid were divided into 
three batches each containing thirty eggs. Each batch was placed 
in 200 c.c. of tap water with 0 5 gram of polished rice. 

‘ A’ was kept in the laboratory at room temperature, i.e., about 
78° F. 

* B * was kept in an incubator at a temperature of about 95 0 F. 
1 C * was kept in an ice chest at a temperature varying from 
56° to 59 0 F. 

For details of the results see Experiment XIII. 


Experiment XIII 




Day 

Catch 

Eff.'S 

3 

4 

5 

6 

7 

8 

9 

10 

1 11 

12 

*3 

H 

15 

* A’ 

30 

iL. 

51- 

31- 

iL. 

2 la. 

12L. 

... 


iP. 

| 2P. 

iP. 

... 

5 P- 

3 M- 

‘B’ 

3 ° 

iL. 



... 

2 la. 

9 l. 


2 L. 


! 

3 l. 

... 

iP. 

w 

‘ C* 

30 

... 



... 


2 la. 

5 l. 

iL. 

... 

iL. 


»L. 

2 L. 

Experiment XIII (< continued) 



Day 

Batch 

Egg* 

l6 

*7 

18 

*9 

20 

22 

23 

21 

2 s 

26 

27 

38 

103 

‘A’ 

30 

IP. 

2M. 

3 E- 

iM. 

3M. 

2M. 

2M. 

2P. 

2P. 

6P. 

iP. 

3 m. 

2M. 

6M. 


... 

* B * 

3 ° 

... 

... 

... 

... 

... 

... 



L. 

w 


... 


... 

• c* 

3 ° 

... 


... 


... 




... j 



ip. 

1.. 

M 


Note :—* L.* = larva (a) pupated and died on 15th day 

' P.’ =* pupa ( b ) the last larva died on 25th day 

1 M.' = adult ( c) this pupa died two days later 

{d) the last larva died on 103rd day 





280 


Thus of thirty eggs kept under the above conditions in the 
laboratory, twenty-four, or 80 per cent., produced larvae, all of 
which pupated and subsequently produced adults. Of the thirty 
kept in the incubator, seventeen, or 56‘6 per cent., produced larvae, 
only one of which pupated ; this one died on the same day. The 
last surviving larva died on the twenty-fifth day. Of the thirty 
eggs placed in the ice chest, fifteen, or 50 per cent., hatched, of 
which number only one pupated. This pupa died two days later. 
The last surviving larva died on the hundred and third day. 

In Experiment XIV, one hundred and ten eggs were divided 
into three batches, ‘ A ’ and * B ’ containing thirty-eight eggs and 
‘ C ’ thirty-four eggs. The methods of storing and the temperature 
were the same as the corresponding batches in Experiment XIII. 
The results are shown in the following statement: — 

Experiment XIV 


I | 

j » Day 


Batch 

Epp j 

5 

6 1 

7 

8 

9 

to ! 

i 

: ; 

11 

! 

1 

12 

! 3 

4 A’ 

3 * 


1 

4 l. 

8L. 

zh. 

iL. 

i 

9 l- 

IP. 


4 B’ 

3 * 

1 

4 L. 

i 



6L. 



8L. ' 


ip. 

(0) 

‘ C* 

34 l 




... 

iL. 


... 1 

1 i 


iL. 

Experiment XIV ( continued ). 


! 




Day 






Batch 


>4 

*5 

16 

*7 

18 

! 9 

20 

21 

47 

4 A * 

3 * 

3 P- 

2P. 

j iM. 

iP. 

2P. 

4M. 

zM. 

■ 

2P. 

iM. 

zM. 


4 B ’ 

3 * 

... 

... 



: 



L. 

w 


‘C* 

34 

... 

! 

12L. 

iL. 

! 

1 

... 


L. 

j£i_ 


(a) This pupa lived only two days. 

(b) The last surviving larva died on ant day. 
(r) The last surviving larva died on 47th day. 





Thus of thirty-eight eggs kept in the laboratory twenty-four 
(63* 1 per cent.) hatched. Of these twenty-four larvae eleven 
(45 8 per cent.) pupated, and ten of the eleven (909 per cent.) 
produced adults. The percentage of adults reared from the thirty- 
eight eggs was therefore 26 3. Of thirty-eight eggs placed in the 
incubator eighteen (47 3 per cent.) hatched, one of which reached the 
pupal stage but survived only two days. Fifteen of the thirty-four 
eggs placed in the ice chest produced larvae, none of which pupated, 
although one lived to the forty-seventh day. The low percentage 
of hatching in ‘A,’ Experiment XIV, as compared with ‘A,’ 
Experiment XIII, is not accounted for. 

The influence of sunlight, diffused light, and darkness on the 
hatching of eggs and development of subsequent stages 

Experiment XV 

In this experiment, one hundred and five eggs were divided into 
three batches, which were treated in the following manner: — 

Batch ‘A,’ containing forty-five eggs placed in 1,300 c.c. tap 
water with a small quantity of fowl faeces, was stored on 
the verandah, where it was exposed to direct sunlight 
during the day and to the lower temperature of night. 

Batch ‘ B,’ containing thirty eggs placed in 1,300 c.c. tap water 
with a small quantity of fowl faeces, was stored in a photo¬ 
graphic dark room. 

Batch ‘ C,’ containing thirty eggs, was treated similarly to ' A ’ 

• and * B,’ excepting that it was stored in the laboratory. 

The results are shown in the tabulated statement XV. 

Experiment XV 




Day 

Batch 

Egg* 

3 

j 

4 

6 

7 

8 

9 

10 

I 1 

12 

13 

* A * exposed to 
direct sunlight 

45 


35 l. 




... 



I2P. 

5 P- 

iM. 

' B * darkroom 

3 o 

... 

24L. 

** *• 


5P. 

| 

! 5 p. 

3P- 

IP. 

8M. 

iM. 

4\i. 

4 C * exposed to 
diffused sunlight 

30 

16L. 


2L. 

2L. 

! 

8P. 

1 

8P. 

13M. 

2P. 

3M. 

2P. 



282 


Expeeiment XV [continued). 






Day 






Batch 

Egg* 


15 ' 16 

17 

18 

*9 

21 

29 

3 2 

4 A * exposed to 
direct sunlight 

45 

2P. 

5M. 

6P. 

2M. 15M. 

iP. 

iM. 

iM. 


iM. 

iP. 

iM. 

‘ B * darkroom 


iM. 

... 1 ... 

1 


iP. 

iM. 

... 



‘ C * exposed to 
diffused sunlight 

3 ° 

i 3M. 

| iM. 


... 

... 

... 




Thus in ‘ A,’ thirty-five (777 per cent.) larvae hatched from forty- 
five eggs, and twenty-seven (77’ 1 per cent.) of these larvae ultimately 
produced adults. 

In ‘ B,’ twenty-four (80 per cent.) larvae hatched from thirty eggs, 
of which number fifteen (62 5 per cent.) produced adults. 

In ‘ C,’ twenty (66'6 per cent.) larvae hatched from thirty eggs, 
all of which larvae produced adults. 

It will be seen that the lowest percentage of hatching occurred in 
the laboratory (Batch ‘ C ’), where 100 per cent, of those which did 
hatch ultimately produced adults. It should be noted, however, 
that no observations were made on the unhatched eggs after the 
thirty-second day ; it is quite possible, therefore, that all, or a large 
proportion of them, may have been naturally resistant. 


2. LARVAE 

The effect of light or absence of light on the development 

of larvae 

Experiment XVI 

Twenty larvae which hatched between 9.45 a.m. and 11.45 a.m. 
on 24th March were divided into two equal batches, and each batch 
placed in a bottle containing 5 or 6 grains of rice and 10 ozs. tap 
water. One bottle (‘A’) was stored in the photographic darkroom, 
the other (‘ B ’) on the laboratory windowsill. All the larvae, 
excepting two of those in ‘A,’ pupated between 5 p.m. on 
27th March and 8.45 on 28th March. On the next day (29th) one 



283 


of the pupae in 4 A 9 died. On the 30th (about 10 a.m.) the 
remaining seven pupae in 4 A 1 and the full number (ten) in 4 B * 
produced adults. 

In this experiment it would appear that the development of the 
larvae under artificial conditions and at room temperature is not 
influenced by the presence or absence of light. 


Experiment XVII. 

4 A/ On verandah exposed to the sun 
4 B.* In dark-room 
4 C.’ In workroom 

4 D.* In ice chest and room temperature 
alternately 


Food given, 

0*3 per cent, in 
1300 c.c. of water. 


Da v. 


h. 

Larvae. 

1 

2 

3 

4 

5 

6 

7 

8 

0 

io 

11 

12 

13 

*4 

i 

*5 

(6 

' I 2 5 

1 








2P. 

5P- 

2 P. 

6P. 

iP. 

2 P. 

iP. 

iP. 











xF. 

iM. 

2 M. 

5 F. 

iM. 

xM. 

2 M 












3F- 

2F. 





' *4 

! 


5 p. 


5 P- 

3P- 

xP. 

A P. 

iP. 

... 




xP. 










6F. 


3F- 















3 m. 

2 F. 

xM. 

4M. 













8 P. 


8 P. 


iP. 

2 P. 


Mi 



... 


j 20 









ioF. 

iF. 


3 F- 

iF. 






1 








3M. 

iM. 





i 



32 










iP. 

xP. 

2P. 

4P- 

2 P. 

2P. 

















iM. 

3 f - 

iF. 


















4M. 


Experiment XVII (continued). 




Day. 

u 

Lanac. 

*7 

18 

>9 

21 

22 

2 3 

24 

2 5 

26 

2 7 

28 

29 

3 o 

3 1 

33 

36 


i 

2 5 


xM. 



... 

... 


... 

xP. 

... 


iM. 

- 




i *» 

xM. 


... 

... 




... 

! ••• 


... 






20 

... 

... 


... 


... 

... 


... 





■ I 



32 

1 

I 

i oP. 
2 F. 

5 P- 

2 F. 

iM. 

IF. 

iM. 

xF. 

xM. 

xM. 


3 F- 

2M. 

i ... 

i 

1 

1 1£ 1 
iM. 

... 

xP. 

... 

! 

iP " 

1 

i \ 

1 

1 

? 


Not/. —* M * = Male. 4 F ' » Female. 4 P * = Pupa. 





284 


Thus in ‘A’ 21 larvae completed their life cycle—84 per cent, 
reared in 28 days. In ‘ B ’ 20 larvae completed their life cycle— 
83 3 per cent, reared in 16 days. In ‘ C ’ all larvae completed their 
life cycle—100 per cent, reared in 12 days. In ‘ D’ 31 larvae 
completed their life cycle—96 per cent, reared in 36 days. 

The effect of temperature and. darkness on the development of 
recently hatched larvae 

The larvae in these two experiments (Experiments XVIII and 
XIX) hatched overnight. In Experiment XVIII, thirty larvae were 
divided into two equal batches, ‘ A ’ and ‘ B,’ each of which was 
placed in 100 c.c. tap water containing 0 25 grams of fowl faeces. 
‘ A ’ was kept in an incubator at 98 6° F. and ‘ B ’ in a second 
incubator which was not artificially heated, the temperature being 
about 78'8°F. The results are shown hereunder:—* 


Experiment XVIII 



1 





Day. 




Batch 

Larvae 

1 

5 

7 

8 

9 

I 

11 

i 

12 13 

14 16 

1 «* 

‘A’ 

>5 

! 

! 

... 

ip. 

« 

... 

L. 

(*) 

... 

... 


. 

1 .. 

! 

4 B ’ 

*5 

1 


\ ••• 

t 

... 

6P. 

6P. 

iM. 

4 m. 

2P. i ... 
4 M. iM. 

iP. 

;M. 

t\i. 


Note. —‘ L * = Larva. ‘ P ’ = Pupa. 4 M ’ = Adult. 

(a) The first larva pupated on the fifth day and died on the seventh. 

(b) The remaining larvae were all dead on the eighth day. 


Thus in A’ (temperature 98 6 0 F.) only one larva pupated, 
and this lived only two days. The remaining larvae died by the 
eighth day. 

In ‘B’ (temperature 788°F.) only 67 per cent, of the larvae 
failed to produce adults. 




285 

Experiment XIX 



! 

Days 

Batch 

Larvae 

1 

10 

1 

II 

12 

13 

14 

*5 

‘A’ 

1 

10 

1 

UA 






‘B* 

10 

3P- 

5P- 

IP. 

IP. 

4 m. 

aM. 

iM. 


(a) All the larvae were dead on the tenth day. 


Thus in ‘A’ (temperature 98'6°F.) none of the larvae survived, 
but in ‘B’ (temperature 78 8° F.) only 10 per cent, of the larvae 
failed to produce adults. 

From these experiments it would appear that excessive heat and 
not darkness was responsible for the mortality. In both experiments 
(XVIII ‘ B ’ and XIX ‘ B ’) the larval period was longer than usual, 
i.e., from ten to thirteen days in the latter. 

Subsequent records of experiments will show that the lengthened 
larval period was probably due to unsuitable feeding, since it has 
been shown in Experiment XVI that the absence of light does not 
retard larval development. 


Quantity and nature of food , and its influence on larval 
development 

In this experiment (XX) forty newly hatched larvae were placed 
in each of five vessels containing 300 c.c. tap water and 01 per cent, 
of food as follows : — 

P.R. 0 3 grm. polished rice. 

M.L. 0 3 grm. dry mango leaf. 

F.F. o‘3 grm. dry fowl faeces. 

B. o'3 c.c. broth, previously exposed to the air for two 

days, with luxuriant growth of bacteria. 

F.M. o'3 grm. fresh fly maggot (dead). 

The average laboratory temperature during the experiment was 
about 7i°F. The results are shown in the corresponding state¬ 
ment (XX). 




286 


Experiment XX 




The mortality totals in the above o' i per cent, foods are 
therefore seen to be:—P.R. 65'o per cent., M.L. 975 per cent., 
F.F. 57 5 per cent., B. 100 per cent., and F.M. 90 per cent. 


Experiment XXI 

The number of larvae used in this experiment varied between ten 
and thirteen to each batch. All were recently hatched. Each batch 
was liberated in a vessel containing 400 c.c. tap water and 
0 25 per cent, of food, as follows : — 


M.L. 

ro gram dry mango leaf. 

D.C. 

ro „ 

dried cockroach. 

H.D. 

ro „ 

dried horse faeces. 

P.R. 

ro ,, 

polished rice. 

G.S. 

2‘0 C.C. 

goat serum. 


The results are shown in the corresponding statement (XXI). 



Experiment XXI 


Batch 

Larvae 

1 

1 

Day 

| 

5 

6 

7 

! 8 

9 

10 

11 

12 

1 

13 

M.L. 

1 

13 

iP. 

iP. 

ip. 

1 

2P. 

1 

IP- 

iP. 

IP. 







3M. 


2M. 

lM. 

iM. 

iM. 

D.C. 

10 


+P. 

2P - 


2P. 






i 




4M. 

1 

2M. 

2M. 




H.D. 

10 





IP. 


2P. 


i p . 








! 


3 M- 


P.R. 

13 

11P. 






• • • 






nivi. 








G.S. 

*3 


ip. 

2 p . 

IP. 

3 P- 



... 


! 

1 

i 




2M. 

zM. 

1 


3 m. 




Experiment XXI ( lontinucd). 


Batch 

Larvae 

Day 



*4 | 

15 

16 

l 7 

.8 

iq i 

20 

21 

M.L. 

*3 


*** 

iP. 


IP. 

iM. 

IP. 

IM. 


iM. 

D.C. 

10 


IP. 


iM. 

... 




H.D. 

10 

IP. 

iM. 


iM. 

... 



... 

P.R. 

13 





iP. 


iM. 

| 


G.S. 

T 3 

2P. 

iP. 

iP. 

! 

3 M- 

iM. 

iM. 

! 



Thus the mortality totals in 0 25 per cent, of the foods shown 
above are as follows:—M.L. 154 per cent., D.C. 10 per cent., 
H.D. 50 per cent., P.R. 77 per cent., and G.S. 77 per cent. 




288 


Experiment XXII 

From nine to seventeen recently hatched larvae were liberated 
in each of nine vessels containing 400 c.c. tap water and 0 3 per cent. 


of food, as follows:— 

M.L. 

i'2gram. dry mango leaf. 

D.C. 

,, dry cockroach. 

H.D. 

,, dry horse faeces. 

P.R. 

,, polished rice. 

P. 

,, peptone. 

P.S. 

0*15 per cent, peptone solution, tweny-i 
filtered. 

B.C. 

1 ,, agar slant, B. coli. 

B.C.2 

1 

2 n * » m j » 

H.U. 

4 c.c. human urine. 


The results are shown in the corresponding statement (XXII). 


Experiment XXII. 


Batch 

Larvae 





Day 





5 

6 

7 

8 

9 

10 

u 

12 

14 

18 

M.L. 

*7 

3 p. 

6P. 


^p. 


iP. 


ip. 







4 \i. 

4M. 


6M. 



2M. 


D.C. 

»7 

2P. 

5 P- 

6P. 


jP- 


••• ' 

... 






iM. 

4 m. 

5M. 


4 m. 1 



H.D. 

9 

2P. 

2P. 





! 







2\i. 



2M. 

i 



P.R. 

i 9 

5 P- 

2 p . 

• • • 


... 

... 

... 


... 







7M. 







P. 

1 

*5 


IP. 

... 


... 

... 

... 


... 

... 

P.S. 

| , 

5 P- 

3 P- 

... 

IP. 


... 


... 






iM. 

3M. 

4 m. 


iM. 





B.C. 

11 

-P- 

2P. 





... 







iM. 

5M. 

2M. 

! 

1 

iM. 


1 


B.C.2 

10 

5 p - 

,p . 




! ... 

iP. 

... 

1 IP. 

... 





5M. 



iM. | 



iM. 

iM. 

H.U. 

9 

4 P- 

... 


IP. 

ip. 


... 


I ‘ P * 





iM. 

2M. 

iM. 

iM. 

iM. j 




iM. 









289 


Thus the mortality totals in O'3 per cent, of the foods shown are 
as follows:—M.L. 5'g per cent., D.C. 59 per cent., H.D. 55-5 
per cent., P.R. 22'2 per cent., P. 93 per cent., P.S. nil, B.C. 18 2 
per cent., B.C.2 20 per cent., and H.U. 22'2 per cent. 

The results of these three experiments are tabulated in Table 7. 

Table 7. 


Batch 

Nature of food 

Percentage of food to 
water 

Percentage of mortality 

P.R. 

Polished rice. . 

o-i 

65 0 



0-25 

77 



0-3 

22-2 

M.L. 

Mango leaf . 

o-i 

97*5 



0-25 

15-4 



0-3 

5’9 

F.F. 

Fowl faeces 

O* I 

57*5 



025 




o -3 


B. 

Broth . . 

0*1 

1 oo-o 



0-25 




o -3 

... 

F.M. 

Fly maggot . 

o-i 

90*0 



0-25 




0-3 


D.C. 

Cockroach 

! 




0-2J 

10-0 



0-3 j 

59 

H.D. 

Horse faeces . 

O-I 




0-25 

5 °-o 



o -3 

55*5 

C.S. 

Goat serum . 

o-i 

• •• 



0-25 

77 



0-3 


P. 

Peptone . 

o-i 




0-25 

... 



o -3 

93*3 

P.S. 

Peptone Solution . 

015 

nil 

B.C. 

1 agar slant B. colt . 

... 

18-2 

B.C.2 

i agar slant B. colt . 


20*0 

H.U. 

Human urine . 

1*0 

22*2 


















290 

Experiment XXIII 

‘A.’ 200 c.c. of tap water containing 05 per cent, rice starch. 

‘ B.’ ,, ,, ,, with the addition of 5 per cent, sugar. 

* C.’ ,, ,, ,, with the addition of half the white of 

an egg. 

‘D.’ ,, ,, ,, containing piece of banana (removed 

after the third day). 

The results are shown in the corresponding tabulated statement 

(XXIII). 


Batch ; Lar/ae 


1 

i 

7 

1 8 

i 9 

1 TO 

I I 

12 

>3 

■4 

*5 

• A* 

| 22 


1 

i 




3 P- 



‘B* 

20 

1 

L. 

fa; 

i 

i 



... 




‘C’ 

20 

... 

j 

qP. 

IP. 

livi. 

6M. 

3 P- 

2M. 

... 

iP. 

3M. 

1 D' 

So 

IP. 

! 8P. 

5 P- 

1 6 P. 

1 

iM. 

4 P- 

4M. 

2M. 

IP. 

9 m. 

2P. 

5M. 



Experiment XXIII ( continued ). 





I 

Batch 

Larvae 

Day 



16 

*7 

l8 

l 9 

21 

22 

23 

24 1 

*5 

4 A ’ 

22 

■ 

JM. 


IlP. 


ip. 

IP. 

11M. 

2 p. ! 

i 

4 M. 

4 B ' 

1 

20 

... 






L. 

(*) 

... 1 

... 

1 

4 C' 

2° 

2 P. 

iM. 


(0 




... 

i 

1 

IP. 

4 D ’ 

3 ° 

... 

6M. 


W 


... 

... 

j 

... 


(«) Some of the larvae now in second skin. 

(b) Larvae in second and third skin. 

( c) Two pupae and one larva died. 

(d) One pupa died. 

(*) The remaining larvae died. 

Mortality ‘A’ 

‘B’ 

‘C’ . 


18 2 per cent. 
100 


35 


10 


i f 





291 


The effects of an increase of temperature on larvae and 
pupae of Stegomyia 

A number of recently hatched larvae were placed in water in 
beakers heated over a water bath. When the temperature reached 
114*8° F., 75 per cent, of the larvae were found to be dead, and the 
remainder failed to develop when the water was gradually reduced 
to room temperature. 

Further experiments were then carried out with older larvae, i.e., 
larvae in their third skin, and with pupae. The procedure was the 
same as in the first experiment. The results of these experiments 
are seen in the tabulated statement (XXIV). 


Experiment XXIV 




Number used 

Number 

subsequently 

reared 

Number killed 
by heating 

Mortality 

i 


T cmpcrsturc 

I.arvae 

Pupae 

I 

90° F. 

10 

3 

all 

nil 

nil 

2 

95° F. 

1 0 

3 

all 

nil 

nil 

3 

ioo° F. 

10 

3 

all 

nil 

nil 

4 

io 5 3 r. 

10 

1 3 

10 

3 larvae 

* 3 % 

5 

no° F. 

10 

j 3 

4 

2 pupae 

7 larvae 

$9'*% 

6 

115“ F. 

10 

1 3 

nil 

| 

100% 


From the foregoing statement it will be seen that the approximate 
maximum temperature that the larvae and pupae are capable of 
withstanding without great mortality is about io 5°F.; beyond this 
temperature there is a very great increase in the mortaliy. Bacot 
showed that a temperature of H2°F. caused a mortality of 
50 per cent., and that higher temperatures caused the death of all 
larvae and pupae experimented with. 

The survival of larvae and pupae out of water 
Experiment XXV 

(1) On filter paper kept slightly moist 

Thirty-four larvae and five pupae were placed on wet filter paper 
and kept moist by adding a few drops of water each day. Two 



adult mosquitoes emerged on the following day, two others on the 
second day and the remaining one on the fourth day. The larvae 
were left on the paper until the fifteenth day, when only two of them 
remained alive. These two were then transferred to a bottle of 
water, but failed to reach the pupal stage. 

(2) On wet filter paper and allowed to 
dry in muslin covered dish 

Larvae and pupae were placed on wet filter paper in a muslin 
covered dish and allowed to dry gradually. Two larvae and one 
pupa were removed to water at the end of thirty-two hours, when 
the paper was nearly dry, and all of them eventually developed into 
adults. 

Of two larvae and one pupa removed at the end of forty-seven 
and a half hours, only the latter was alive, but this died soon after 
in a dish of water. Another pupa, however, was alive after fifty and 
a half hours on the paper, but was dead at seventy-two hours. None 
of the larvae appeared to have survived beyond thirty-two hours. 

(3) On wet filter paper and allowed to dry 
in a glass covered dish 

Thirty-one larvae were placed on wet filter paper and put on 
a glass covered petri dish. After sixty-five hours in this dish seven 
larvae were removed to water, where one of the seven eventually 
matured. The remaining twenty-four larvae were transferred to 
water after having been on the paper for eighty-nine hours. Of 
these five were alive, but one died during the following twenty-four 
hours. The remaining four lived for nine days, after which no 
further record of them was kept. 

3. ADULTS 

l.ength of life of adult mosquitoes in captivity when unfed 
and fed on various foods 

The mosquitoes used in this experiment (XXVI) were recently 
emerged laboratory-bred insects, all of which (except A.) were given 
an opportunity of feeding daily during the course of the experiment. 
The various batches were kept in similar cages and treated similarly 



2 93 


to each other, excepting with regard to the food given. The 
average daily temperature was between 7 i°F. and 8o°F. 

A. Not fed. 

B. Fed on syrup. 

C. Milk and sugar. 

D. Banana. 

E. Blood once (females); banana afterwards. 

F. Blood for sixty-one days, then bananas (females); banana 

only for males. 

The results are shown in the following tabulated statement: — 


Experiment XXVI 


E\pt. 

! 

Number 
of adults 
used 







Day 




2 | 

4 

7 *2 

*5 

! 9 

26 

3 ° 

35 

17 

5 * 

5* 

61 

68 

93 

A ’ 

! 

*;F. 

qF. 

>?F. 

iF. 










1 



IZM. 

XVf. 

4 m. 






1 






B’ 

?F. 



2F. 

iF. 

2F. 

iF. 


... 1 



1 




4 M. 


2M. 



iM. 


iM. 




1 



r * 

7 F- 



iF. iF. 

2F. 

2F. 





... 

... ! ... 




2\I. 



1 2 M. 









i 



) * | 

CF. 





iF. 


iF. 

iF. 

iF. 



! ... ! iF. 



5M. 





iM. 

iM. 

iM. 

iM. 

iM. 



j 1 


;• 

3 F. 





... 

iF. 


iF. 

... 

... 


if. ; ... 1 ... 


iM. 



1 



M 

iM. 




1 1 1 

• 

15F. 



... 5 F. 


5 F. 


iF. 

... 

iF. 

iF. 

iF. 


i iF. 


5M. 



, 


iM. 


2M. 


iM. 



iM. 

1 


(r) — Escaped from cage. F. = Females. M. = Males. 


Rrtention of eggs by gravid females 
Oviposition on dry surfaces 

It is a generally accepted belief that either water or a damp 
surface is essential to oviposition, and that if such is not available 
the gravid female will retain her eggs for a long period—until her 
death or until favourable conditions are made available. My 
experience in the routine of breeding Slegomyia , as well as the 
results of a number of experiments undertaken especially to confirm 
former observations, convince me that this belief is well founded. 



294 


Oviposition on oily surface 

That similar results follow the provision in the breeding cages of 
only water having a surface film of oil is shown by the following 
experiments: — 

Experiment XXVII 

On the morning of lith September, two gravid females and four 
males were placed in a breeding cage containing a dish of tap water 
covered with a thin film of kerosene. For several days previously 
the females had fed on human blood and the males on banana, which 
foods were offered daily during the progress of the experiment. The 
following observations were subsequently recorded : — 

nth September (afternoon), one female fed. 13th, one female 
fed. 14th. one female fed; three males drowned. 17th, a female 
drowned. 19th, the remaining male drowned, three others liberated 
in cage. The remaining female fed on 29th September and on 
4th October. No eggs laid up to this date. A dish of tap water 
without oil on surface was put into the cage, and the first dish (with 
oil) removed. 5th October, forty-six eggs laid over-night, the first 
oviposition during her twenty-four days of captivity. Female fed in 
afternoon. 15th October, seventy-nine eggs laid over-night. Dish of 
fresh water removed and replaced by one containing water with oily 
film. 16th October, female fed. 22nd October, female fed, and at 
intervals of about three days until her death. One male drowned. 
24th October, one male drowned. 25th October, one male drowned. 
8th November, female drowned. 

Egg laying by unfertilised females 

We have noticed on several occasions that certain batches of eggs 
produced females only, and that when segregated and fed on human 
blood these unfertilised insects were capable of laying eggs. 

In this connection the following experiment is of interest as 
showing the increase which takes place in egg production after 
fertilisation. 

The mosquitoes used in this experiment, seven in number, were 
reared from a batch of eggs which produced females only, and as soon 
as possible after hatching they were segregated in such manner as to 
preclude the possibility of fertilization. Opportunities for feeding 



2 95 

on human blood were given frequently from the date of emergence, 
i.e., 7th September. After thirty days of captivity in this manner, 
two males were liberated in the cage with the five surviving females 
for seventeen hours, with the results shown in Experiment XXVIII. 

Experiment XXVIII 


Dates on which 
eggs were laid 

Number 
of eggs 
laid 

Approx. 

number 

proved 

fertile 

Fertility 

Deaths to 
date 

Feeds 
given to 
date 

Remark? 

22.9.17 

5 


1 

2 

14 


24.9.17 

6 




16 


1.10.17 

85 

i 



l 9 

Total number of 
eggs laid by 
unfertilised 
females : 105 

8.10.17 

9 

i 



23 

Total number of 
feeds given : 23 


4 p.m. 8.10.17 ) 

to - (17 hours) 2 males liberated in cage with 5 surviving females. 

9 a.m. 9.10.17 ) 


10.10.17 

60 

30 

+ 


28 


12-13.10.17 

7 8 

7 « 

+ 




15.10.17 

12 

3 



3 i 


17.10.17 

3 1 

3 1 

+ 


33 


18.10.17 

60 

60 

+ 


35 


22.10.17 

140 

140 

+ 


39 


25.10.17 


38 

+ 

1 

41 


20.10.17 

120 

100 

+ 

... 



2.11.17 

72 

60 

+ 

1 



7 * 1 I * I 7 

140 

100 

+ 

1 

44 

1 

Total number of 
eggs laid by 
fertilised 
, females : 765 

8.11.17 




2 


Total number of 
feeds given to 
fertilised 
females • 21 


From the foregoing it will be seen that seven unfertilised females 
did not lav eggs during the first fifteen days, and that during this 





296 


period two died. During the following fifteen days the remaining 
five females laid one hundred and five infertile eggs and were then 
mated with two males. During the thirty days following the intro¬ 
duction of the males seven hundred and sixty-five eggs were laid, of 
which number approximately six hundred and forty proved fertile. 


Experiments to ascertain whether Stegomyia fasciata 
ivould lay eggs when fed on food other than blood 

From time to time various batches of mosquitoes were given food 
other than blood for periods ranging from twenty-five days to 
one hundred and forty days. The following foods were experi¬ 
mented with : — 


Concentrated sugar solutions 
Concentrated peptone and sugar solution 

r_ j.___1 1_11: 


solution 

Milk and sugar 
Banana 

Peptone solution 

Syrup 

Honey 

Dates 

Apple 





3 

f * 




2 

t y 




1 

>) 




2 

» j 




1 

j 1 

... 



1 

1 j 




1 

t > 




1 

f 1 


In each of the three experiments in which peptone and sugar were 
given as food a number of eggs were laid, about 60 per cent, of 
which were fertile. In all other experiments there was no egg 
production. 


ACKNOWLEDGMENT 

In conclusion, I desire to thank Mr. G. F. Hill, Entomologist to 
the Institute, for his help and advice in the preparation of the 
manuscript of this paper for publication. 














297 

ANCYLOSTOMA CEYLANICUM 
IN THE CAT IN DURBAN 

BY 

B. BLACKLOCK 

(Received for publication I August , 1919) 

Material from Durban was sent by Dr. F. G. Cawston on 
10th June, 1919, with the following notes: — 

1 Ankylostoma duodenale from a cat in Durban. The gut also 
contained a tapeworm, and death seems to have been caused by the 
numerous Ankylostoma/ 

The material consisted of ten female worms, of which nine were 
complete and in a good state of preservation. 

It is not possible in the absence of specimens to describe the 
characters of the male bursa, but the general appearance of the 
females, the measurements and the arrangement of the chitinous 
buccal capsule and teeth have been studied. 

Of the nine complete specimens the minimum length is 7 0 mm., 
the maximum 9*5 mm., the average 8*3 mm. The arrangement of 
the teeth is characteristic. There is a pair of large ventral teeth 
and also a very small pair situated at the base of these on a deeper 
plane. The measurements and mouth structure are, in fact, such 
that this parasite cannot be distinguished from A. ceylanicurn 
(Looss). This parasite was recorded by Yorke and Blacklock 
(1915) in seven dogs in Sierra Leone, and was further referred to 
by them (1917) subsequent to Macfie’s discovery of the same 
parasite in four of ten dogs at Accra (1916). 

That this parasite should be found in Durban is of interest, 
more especially so if, as Dr. Cawston surmises, it should be the 
cause there of fatal disease in cats. 

REFERENCES 

Macfie, J. W. S. (1916). Report of the Accra Laboratory for the year 1916. 

Yorke and Blacklock, (1915). Ankylostomiasis in Dogs in Sierra Leone. Attn. Trop. 

Med. & Parasitol ., Vol. IX, p. 425. 

-( I 9 I 7). The Occurrence of Ankylostoma ceylanicurn in West African Dogs. Ann. 

Trop. Med. & Parasitol ., Vol. XI, p. 69. 




THE 

MECHANISM OF THE SPONTANEOUS 
ELIMINATION OF YELLOW FEVER 
FROM ENDEMIC CENTRES 

•BY 

H. R. CARTER 

ASSISTANT SURGEON-GENERAL U.S. PUBLIC HEALTH SERVICE 


(Received for publication i October , 1919) 

First: This inquiry is limited to places in regions in which 
yellow fever is endemic: biologically, to places in which Stegomyia 
is abundant and active at all seasons: geographically, to tropical 
America. 

The mechanism of its spontaneous elimination from places north 
or south of this endemic zone is too well known to be discussed, 
viz., the death or inactivity of Stegomyia caused by cold weather. 

Secondly: It is limited to places in the above regions in which 
elimination was not due to the sanitary measures taken for this 
purpose. 

The mechanism by which this has been accomplished is in effect 
the same as I have just mentioned as occurring north and south of 
the endemic zone, i.e., the control of the insect host. It differs from 
the elimination due to cold weather only by the degree of destruction 
of Stegomyia and the method of accomplishing this. Indeed, in 
places far south—as in Key West—the degree of the elimination of 
Stegomyia by the advent of winter may not exceed that of a well- 
conducted campaign against their breeding-places in Panama. 

Note that in such towns it is not necessary to exterminate 
Stegomyia to eliminate yellow fever. If the number of mosquitoes 
be brought below the ‘critical number’ for yellow fever—an idea 
we owe to Ross as applied to malaria—at that place the disease will 
die out. Note, too, that this ‘critical number’ for any place will 
vary directly as the proportion of men immune to yellow fever to 
the total population : thus, if with one hundred cases of yellow fever 



300 


introduced into a community in which all were susceptible to yellow 
fever, the number of Stegomyia were such that exactly one hundred 
men would be infected from them, the disease would neither die out 
nor increase. This would be the critical number of Stegomyia for 
that place and time. With less mosquitoes than this, less than 
one hundred men would be infected and the fever would die out. 
If more, it would increase. Now if one-fourth of the inhabitants 
are immune to yellow fever, obviously the same number of mosquitoes 
which infected one hundred men before would now infect only 
seventy-five—one-fourth of their bites going to immunes, and hence 
wasted—and the disease would die out. The number of mosquitoes 
required to infect one hundred more men, and hence perpetuate the 
fever, would have to be increased by one-third above the first 
number. 

Obviously then, this critical number, below which the Stegomyia 
must be brought to eliminate fever, is less in a town as the proportion 
of susceptible people increases, and more intensive work is required 
to eliminate it from such a community, other things being equal, 
than from one in which a large proportion of the population is 
immune. This will be referred to later. 

Yellow fever has been driven from the great permanent endemic 
foci of Havana, Vera Cruz, Panama, Rio, Santos and Para by 
sanitary measures, i.e., by control (lessening) of Stegomyia. It has 
also disappeared from a number of former endemic foci in which no 
measures were taken and in which Stegomyia are still abundant and 
active. This is so well known that it only requires mention. 
Among them I name Georgetown and Demarara in British Guiana— 
the former the seat of the several epidemics on which Blair’s 
invaluable monographs are based—free now as are Kingston and 
Port Royal in Jamaica. Add Port-au-Prince, Jacmel, Cape Haytien 
and San Domingo, in none of which can sanitation have been a 
factor, for there had been none. Yellow fever was virulent in all 
four for years, and to the last named is accredited the evil 
distinction of being the place where this disease was first introduced 
to the white race. St. Thomas, accounted a danger to naval vessels 
for many years as always infected, has been free for over twenty-five 
years; as has St. Lucia, the site of severe epidemics, except for one 
epidemic in 1901 from infection introduced from Brazil (Low). 



3oi 


Except St. Lucia none of these have, I think, reported yellow fever 
for from twenty to thirty years;. 

Havana was freed from yellow fever by sanitary measures— 
control of Stegomyia breeding. Nothing of the sort was done at 
Puerto Principe, where there was a sharp epidemic in 1899. I think 
the same is true of Santiago de Cuba, with an epidemic the same 
year, but I have less knowledge of the sanitary measures taken 
there. Cienfuegos and Matanzas, to my positive knowledge, were 
free from yellow fever long before this sanitary measure was 
instituted for them. All four of these Cuban towns are free now, 
and have long been free. La Guira—once of evil fame—Maracaibo, 
Cartagena and Corinto, all formerly infected, have shown no fever 
for a number of years. I could add many others. 

What is the status of these towns as regards yellow fever ? Has 
the disease simply ceased to appear, being still existent, or has it, 
that is the parasite, ceased to exist in this community ? In other 
words, does the ‘ Spontaneous Elimination,’ of which I have 
purposed to show the ‘ Mechanism,’ occur ? This really is the first 
proposition to be proven. 

The reappearance of fever in certain towns after periods during 
which it has not been reported, has made us, especially those of us 
charged with Maritime Quarantine, slow to accept a place, especially 
a seaport, as free from yellow fever simply because none has been 
reported there for some time; and this without impugning the good 
faith of the local health authorities. Yellow fever can exist among 
the children of a town, especially if there be much negro blood 
among the people, and be scarcely capable of diagnosis. It would 
practically not be reported among such. If on-report of yellow fever 
does not of itself imply its non-existence. 

We need not consider the old theory that although it is not 
appearing in men, yet the organism is still alive and growing in 
fomites ready to attack any susceptible man who is exposed to it. 
And yet I am persuaded that part of this idea, that of the 
permanence of the condition, is the parent of the concept of * latent 
yellow fever.’ That it is carried on indefinitely ‘ in endemic 
centres ’ by ‘ recurrent attacks among the indigenes ’ is the doctrine 
of a Commission of the Pasteur Institute. The r 61 e of human 
carriers without symptoms in thus continuing the disease would be 



added by some writers. This has been further modified by the 
doctrine of ‘ larval yellow fever *: that the strain has been so 
attenuated by passing through the resistant indigenes that the 
disease is taken lightly even by newcomers (Europeans) and hence 
not recognized, although immunizing them. In any case the general 
belief has been that a town or district infected with yellow fever in 
the tropics rarely frees itself, that is without sanitation, from the 
infection. 

It may not be easy to pronounce such a town free from yellow 
fever. Have we then no test to determine the presence of. yellow 
fever, the existence of the parasite, in a community ? The positive 
test is simple and convincing. If a case of yellow fever is contracted 
in that community, yellow fever exists there. The negative must 
be that people suceptible to yellow fever, and in whom its diagnosis 
would be made if they developed it, live in this community so as to 
be exposed to yellow fever if it exists therein and are not found to 
develop it. 

Negative evidence is convincing in proportion to its mass, and 
to accept this negative test as convincing we must have large 
numbers of exposures of such people, i.e., many people exposed over 
a considerable period of time, and the more intimate the local 
relations with the native population among which the fever is 
supposed to be latent the better. 

So far as ‘ larval yellow fever ’ is concerned, the advocates of this 
phase of the disease admit, or rather assert, that the passage of 
the organism through newcomers restores its virulence and the 
disease becomes recognizable. Prolonged residence then, of large 
numbers of newcomers in the infected environment should do away 
with this camouflage of the disease, for the existence of which, 
indeed, I have seen no satisfactory evidence. Is not this test fair ? 
Let us apply it. I think it will be admitted without argument that 
Georgetown and Demarara, Kingston and Port Royal are now free 
from yellow fever. We received men at Ancon who had contracted 
yellow fever in Corinto in 1905 and 1906, when it was also at 
Managua and Valencia. We kept an Inspector at Corinto from 
1908 to 1912, a man who had not had yellow fever. He reported 
no yellow fever, no sanitary measures taken and Stegomyia in 
abundance. In the spring of 1912, five thousand U.S. Marines 



3°3 


landed at Corinto, making this place their base of operations. They 
occupied this place and Managua and Valencia all summer. No 
yellow fever occurred among them. Could there have been any in 
Corinto? We have had Marines at Jacmel, at Cape Haytien, at 
Port-au-Prince and at San Domingo. Prior to this, and when we 
had no control of port sanitation, we had naval vessels lying at these 
ports and landing parties and no yellow fever among either garrisons, 
crews or landing parties. Yet in 1905, of two naval vessels lying 
in Panama Bay, Panama being infected, one became infected— 
seven cases and two deaths—although they took all the precautions 
their regulations called for, certainly more than were possible in the 
close harbour of Port-au-Prince or when sending armed men ashore. 
So in Guayaquil, in 1907, I think, one naval vessel.lay in the 
harbour and she developed yellow fever aboard, losing her 
Commander. All of these San Domingan towns had had yellow 
fever badly and for many years, infecting the vessels, naval and 
others, in their harbours. 

In Maracaibo, in 1916, we found Americans working the oil 
wells. There was a large number of them, scattered all over the 
town; they were practically all susceptible to yellow fever and they 
had been there some eighteen months or over. Maracaibo was alive 
with Siegomyia. There had been no yellow fever among the 
Americans. Obviously there was none in Maracaibo. 

Baranquilla is the Caribbean port of entry to the Colombian 
highland. Through it pass each year some thousands of people 
susceptible to yellow fever: Colombians from the plateau, Americans 
and Europeans. These people, going either way, wait in town 
from one to six days for vessels. No yellow fever has been reported 
among them since 1907 or 1908. It could scarcely have been missed. 
Guiteras and myself in 1916 made a most careful examination of the 
mortuary records of the town since 1911, and we are sure that 
yellow fever has not been prevalent among the children since that 
date. Baranquilla is innocent of any yellow fever sanitation. If 
Baranquilla stays free, Cartagena is free also. 

Why multiply examples ? I could give a number of others, but 
I think I have shown that by our test strictly applied towns in the 
tropics do free themselves from yellow fever, and that without 
sanitary work. This is the first part of my proposition. The 
examples I have given show that this is not rare. 



3^4 


By what mechanism is yellow fever thus eliminated ? 

If we assume that one attack of yellow fever gives, usually, a 
permanent immunity to that disease the argument will be easier to 
follow. I, myself, believe that this immunity is generally permanent, 
as much so as that from small-pox or measles. Other men do not. 
It will be seen that this assumption of permanence is not essential 
to the argument for the mechanism presented. 

From the known facts of the conveyance of yellow fever, it is 
obvious that the conditions for the continued existence of yellow 
fever in a community are three : the carriers, active Stegomyia 
calofus and susceptible men: all present at the same time, the 
insects having access to both classes of men. 

Parasites exist only in an infected mosquito or in an infected 
man. They live in the mosquito only during its life, and only a 
short time, infective to mosquitoes, in man. 

Here, then, are two postulates: — 

(1) Since the parasites in the mosquito live only during the life 
of the host, say, ten days, no interval greater than ten days may 
elapse between the date when some sick man infective to mosquitoes 
is bitten by them and the date when one of the mosquitoes infected 
by him feeds on a man susceptible to yellow fever without the death 
of the parasites, and hence the extinction of yellow fever in that 
community. 

(2) Susceptible people, then, are necessary for the continuance of 
yellow fever in a community. Such people must not only be 
present, but must be present under certain conditions of time and 
place with relation to the Stegomyia infected from other people with 
yellow fever. If in a community there be no susceptible people 
fulfilling these conditions, yellow fever will disappear. 

Now let us consider a community in the tropics in which yellow 
fever is present, Stegomyia abundant and active at all seasons, and 
with susceptible people. Parasites, of course, are present in those 
sick of yellow fever, and, since Stegomyia are active all the year 
round, this place will be an endemic focus of yellow fever. 

Obviously, if one attack of yellow fever produces in general a 
permanent immunity, such a community will have in time no people 
susceptible to yellow fever left, unless there is an introduction of 
such people. Yellow fever would then disappear, and, as soon as 



3°S 


the infected mosquitoes died off (within our ten days), the parasites 
would disappear and the community be free from infection. 
Indeed, yellow fever would doubtless disappear before there were 
‘ no people susceptible to yellow fever left,’ because, under the 
doctrine of chances, there would be no susceptible people left 
fulfilling the conditions of time and place mentioned above before 
there were absolutely hone at all, possibly long before. Once free, 
it would remain free for ever, unless the same three factors for 
conveyance are again brought together. In the natural course of 
events a new generation would grow up susceptible to yellow fever, 
susceptible immigrants move in, and Stegomyia breed to the limit; 
but unless the parasite be again introduced the community would 
remain free from yellow fever. In such a community, growing 
naturally, an epidemic would result if the parasites were introduced 
some years after it had been free of infection, the maximum age of 
the natives then developing the yellow fever depending on the length 
of the interval of freedom. 

An immigration of susceptible people, then, is necessary for the 
continuation of yellow fever in a community, and if this immigra¬ 
tion fails, or fails to fulfil certain conditions, yellow fever 
disappears. This mechanism I have. called the ‘ elimination of 
yellow fever by failure of the human host.’ 

This immigration might be of susceptible people from some other 
place, or of infants bom in the place itself. This is just as true an 
introduction of susceptible people as the other. As old Blair says, 
‘ truly are they new-comers.’ If these additions to the susceptible 
population conjointly fulfil the conditions necessary for the 
continuance of yellow fever, as I have stated them, it will continue; 
if they do not, it will disappear. 

The effect in continuing yellow fever of each class of these 
additions to the susceptible population—men from outside the 
community, and babies born in it—will depend on many factors, 
but among others on its amount, increasing for each class as that 
class increases. 

Both classes of immigration then affect the continuance of yellow 
fever, and theoretically either one may be sufficient to continue it. 
Yet the proportional effect of the introduction of an adult and the 
birth of a baby in keeping up the infection is very different, that 



306 


of an adult immigrant being very much the greater; so that to 
supply the susceptible people necessary to keep up yellow fever, it 
requires a very much larger number of babies to be bom than of 
susceptible immigrants. Adult immigrants are of far more 
importance in thus keeping up yellow fever in proportion to their 
numbers than babies, and a town receiving no susceptible immigra¬ 
tion needs to be mjich larger to be a permanent focus of yellow 
fever than if it did receive such immigration. Gorgas, indeed, in 
1916 expressed himself as believing that immigration of susceptible 
adults was necessary to continue yellow fever, that it could not be 
kept up by the births alone. I cannot think so. 

How does our explanation agree with the known epidemiology 
of yellow fever and with the facts we have adduced in our first 
proposition? On this agreement will depend whether we hold it 
tenable or net. Obviously, if our explanation holds, it will be the 
small and moderate sized towns receiving little immigration which 
will free themselves of yellow fever; the large cities and those with 
much susceptible immigration not doing so. And so we find: 

(1) The towns I have mentioned as freeing themselves of yellow 
fever are all small, or of moderate size, and out of the way of 
commerce, receiving little immigration of suceptible people, and this 
phenomenon is evidently common among such towns. 

(2) It is the large towns and those with considerable susceptible 
immigration: Guayaquil, Havana, Vera Cruz, Rio, etc., which did 
not free themselves, or are not yet free, from yellow fever. 

(3) Besides these there are certain communities of small towns 
between which the travel relations are so close that they must be 
held, for this mechanism, as a single large town. Indeed, in such 
a group of inter-related towns the propagation of yellow fever would 
be decidedly slower than in a single town of their joint population, 
and the fever would therefore last longer and be less apt to disappear 
than in the single larger town. Such a section is Yucatan and 
Campeche—the large sisal haciendas being indeed towns-r-and this 
section has not freed itself from yellow fever, but seems to be a 
permanent endemic focus of yellow fever needing sanitation for its 
elimination. 

(4) Also, towns which were badly infected while prosperous— 
hence large and with much immigration—as Georgetown and 



3<>7 


Demarara, Port-au-Prince and Cartagena, become free from yellow 
fever as their trade declines. And see how Guayaquil reverses this! 

(5) Also we find these small towns liable to epidemics of yellow 
fever from time to time, when a sufficient number of susceptible 
people have accumulated and the parasite is again introduced, and 
that when this occurs, as at Buenaventura in 1915 and 1916, 
‘children and people who have moved in during the past ten years 
were attacked.’ The last epidemic here was twelve years previously. 
The occurrence of epidemics among the native-born is, to me, proof 
that yellow fever had not been general, whether ‘latent ’ or not, since 
they were, say, three or four years old. 

It is this recurrence of fever which has given rise to the belief in 
‘latent’ or ‘larval’ yellow fever held by many very eminent men. 
Unquestionably yellow fever may, and at times does, exist 
unrecognized among the native children of a community; showing 
itself only, or rather being recognized only, when it attacks some 
stranger. Here we have true recrudescence whenever an influx of 
strangers occurs. 

This view is too well known to require elaboration. It is true, 
and I will not pretend to predicate how long such a condition 
may last; nor deny that, under some conditions, it may last 
indefinitely and by this means alone keep the place or area of 
communicating places a permanent focus of yellow fever, and 
without cases ocfcurring sufficiently marked to compel recognition. 
What I do deny is that this is the rule. Indeed, I believe that it is 
rather the rule for yellow fever to disappear, to disappear completely, 
from isolated communities of moderate size and this without sanitary 
work or diminution of Stegomyia. 

The instances given prove that the spontaneous disappearance of 
yellow fever is not rare. An analysis of them would show that, in 
the absence of adult immigration and of inter-travel, this is to be 
expected to occur in a large proportion of towns, and that after this, 
even when there are influxes of strangers, outbreaks in such towns 
do not occur, unless they are in communication with some 
infected focus. These outbreaks, .then, are re-infections and not 
recrudescences of ‘ latent ’ or ‘ larval ’ yellow fever. 

I think our explanation then agrees, and agrees completely, with 
the facts as we have found them, and with the known epidemiology 



308 


of yellow fever. Not only is it consistent with the facts observed, 
but the deductions from it are in accordance with other facts not 
hitherto noted, or at least not stressed. On this kind of evidence 
we are accustomed to accept other laws of nature. 

It is obvious that the explanation I have given of the spontaneous 
elimination of yellow fever depends absolutely on the doctrine that 
an attack of yellow fever confers immunity against another attack. 
In proportion as that immunity is permanent and general, the chance 
of the exhaustion of the susceptible human material by a definite 
number of cases of fever, that is at a definite time, to the point of 
causing the disappearance of disease is greater than if the immunity 
be of short duration and uncertain. If this immunity be not 
permanent, but yet endures for some time, the disappearance of 
yellow fever by the mechanism I have outlined can still occur. This 
is evident. It is also evident that the men in our community who 
lose their immunity through lapse of time add to the supply of 
susceptible material just as immigrants would. In proportion as 
attacks recur frequently and at short intervals, so will the chance of 
failure of the human host, to the point of causing disappearance of 
the disease, diminish. If ‘frequently recurring attacks of the 
indigenes * are the rule, and these recurrences are indefinite, they 
might very well continue the fever in a very mild or ' larval 9 form 
indefinitely, independently of immigration or new births. The 
possibility of disappearance by the mechanism we have given is then 
not dependent upon the permanence of the immunity given by one 
attack, but the chance of its occurring in any place at any definite 
time is directly dependent on it and reaches its maximum when, as 
we believe, one attack gives permanent immunity. The frequency 
with which yellow fever has disappeared when immigration was 
lacking is evidence against recurrent attacks. 

Similarly, it fails in the presence of a permanent reservoir host 
for the micro-organism of yellow fever infective to Stegomyia or 
otherwise capable of communicating it to man, analogous to those 
for the trypanosomes of sleeping sickness. The existence of animal 
hosts, other than man, with the same reactions to the parasite as 
man—i.e., in which an immunity was caused by an infection, 
temporary or permanent—would not seriously affect it. Such 
animals, if present, would simply count as a certain extra number 
of men. 



3°9 


The evidence for the existence of a permanent reservoir host 
seems to me to have no basis in observation. The evidence against 
it, or rather against it being an animal associated with man in the 
places in which yellow fever has been studied, of necessity negative, 
is considerable. 

Nevertheless, it would be much easier to accept the American 
origin of yellow fever, for which the historical evidence is very 
strong, if we had evidence of the existence of such a host in 
Columbian and pre-Columbian times in the Antilles or on the 
Carribbean litoral. 

One thing we must note, however: The fact that yellow fever does 
disappear from towns in the tropics without sanitary work and with 
Stegomyia abundant is true. The explanation I have given is 
logical: it is in accord with what we know of the epidemiology of 
yellow fever, and I believe it to be true. Nevertheless, it is only a 
deduction from observed facts, not the fact itself. 

One other thing. For a town which has freed itself from yellow 
fever by the failure of the human host to remain permanently free 
from yellow fever, isolation from infected places is necessary. When 
yellow fever has been eliminated by the control of the insect host, 
this isolation is not necessary as long as this control continues to be 
efficient, because with the control of the insect host, yellow fever is 
not communicable, and such parasites as are brought in by infected 
men or infected mosquitoes would, at the most, establish a very 
temporary focus of infection. If the control were complete, infected 
men would transmit no parasites. It is to be noted, however, that 
the reduction of Stegomyia sufficient to eliminate yellow fever from 
a town in the tropics would nearly always be less than that required 
some years later to prevent its spread, because there will then be a 
larger proportion of susceptible people than at first. Hence the 
mosquito control must be more intensive. 

It is worth contrasting these two methods for the elimination of 
yellow fever. That by the control of the insect host is unquestion¬ 
ably the method of election for the sanitarian, while the method by 
which it is eliminated in nature is by failure (control) of the human 
host. 

For malaria the method by which it has been eliminated 
in nature is by control of the insect host (agricultural drainage). 



3io 


Malaria confers minimal immunity, and human ‘carriers’ without 
symptoms are in effect reservoir hosts. 

Every one has noted the great diminution of yellow fever which 
has been going on since the fall of the tropical sugar industry in the 
smaller West Indian Islands, Hayti and the Guianas, and which is 
still in progress. It is due to a variety of causes: -— 

(1) To the diminished trade and commercial importance of the 
Islands, and the Carribbean litoral, and hence diminished immigra¬ 
tion to this region. 

(2) The substitution of steam for sailing vessels has enormously 
lessened the number of infections (parasites) carried between ports, 
thus lessening the re-infection of such ports as had cleared them¬ 
selves of yellow fever. Sailing vessels frequently carried Stegomyia 
breeding in their water supply, as well as infected men, while iron 
steam vessels very rarely carried the former, and hence, in spite of 
the shortened voyage, were much less efficient in the transport of 
parasites. Sailing vessels, too, lay longer in port, had larger crews 
per unit of carrying capacity and were under laxer discipline, thus 
furnishing a much larger number of susceptible men ashore— 
temporary immigrants—in the ports they visited. 

(3) With the loss of its commercial importance, indeed before it, 
came the diminished strategic importance of the Carribbean Sea and 
the practical withdrawal of European fleets and garrisons. In the 
18th and the early part of the 19th centuries, the Carribbean was the 
rendezvous of the fleets of Great Britain, France and Spain, and to 
a certain extent, of the United States. The war vessels, sailing 
ships then, lay long in port; some in permanence as receiving ships 
for crews fresh brought from Europe. They carried large crews, 
one thousand men or more, and their shore parties added markedly 
to the susceptible population of the Carribbean seaports. They 
were a prime factor in carrying yellow fever from port to port both 
in infected men and in infected mosquitoes. It is worth noting how 
very generally the introduction of yellow fever in the different ports 
is ascribed to a naval vessel. 

The r 61 e of the European garrison, with its frequent drafts from 
Europe and its massing of susceptible men, was the same in 
continuing the infection at the garrisoned port. 

(4) The extinction of the great permanent foci of Havana, Vera 



3 11 


Cruz,. Panama and Rio. With this a number of less important places 
were freed from yellow fever by sanitary work in control of the insect 
host; Santos, Para, Manaos,. Iquitos, Pernambuco and Caracas. 
This enabled such ports, on the Carribbean especially, as could 
spontaneously free themselves of yellow fever to remain free, being 
no longer exposed to infection, or rather much less so exposed. The 
smaller ports, freeing themselves from yellow fever from time to time 
by failure of susceptible people, had been doubtless continually 
re-infected from the permanent foci, whenever they had accumulated 
sufficient susceptible material, by birth or immigration, for the spread 
of the fever. Even the Great War has helped in this, stopping 
European immigration, and by the commercial depression it caused 
lessening travel in the American countries in which yellow fever 
prevailed. To it is probably due the freeing of Maracaibo, Cucuta 
and Bucaramanga. 

It is facts like these which justify the plan of the International 
Health Board for the elimination of yellow fever from the earth, a 
plan already operative at Coro and at Guayaquil. In my judgment 
it is entirely feasible. Many places in which yellow fever exists will 
need but minimal sanitary work to turn the scale against it, and the 
freeing of one place from yellow fever frequently prevents the 
infection of some other, making an endless chain of good. Many 
of tl^e higher forms of life have permanently disappeared from the 
earth, some in our own times, but this is the first attempt made for 
this purpose againSt a micro-organism pathogenic to man. Its 
accomplishment will mark an era in sanitation. 




3»3 


THE METABOLISM OF WHITE RACES 
LIVING IN THE TROPICS 

III. THE INFLUENCE OF EXTERNAL 
TEMPERATURE AND RATE OF COOLING 
UPON THE RESPIRATORY METABOLISM 

BY 

W. J. YOUNG 

(Received for publication 13 October , 1919) 

From the Biochemical Laboratory of the Australian Institute 
. of Tropical Medicine 

A considerable amount of attention has been devoted to the 
question whether the human organism responds to changes in 
external temperature by changes in the quantity of heat produced; 
that is to say, whether changes in the heat production play any part 
in maintaining a constant body temperature. The action of low 
temperature has been studied by Loewy (1890), Johansson (1896), 
Rubner and Lewaschew (1897), and others, by observing the 
respiratory metabolism under different conditions, and the results 
point to the conclusion that an increased metabolism is only 
produced when the cold is sufficient to stimulate the muscles to 
voluntary or involuntary action, and to' cause shivering. The 
increased metabolism produced on a cold day is due, therefore, to 
the cold air acting on the skin and producing an increased muscular 
activity or tone. 

With regard to high external temperatures, such as exist in the 
tropics, the question is of considerable interest as to whether the 
body adapts. itself to the altered conditions by a decrease in its 
heat production, or whether the temperature is regulated entirely by 
heat loss, i.e., by physical means. 

A summary of the work done on the subject of metabolism in 
the tropics from this point of view has been given by Breinl and 
Young (1919) in an article on the settlement of Tropical Australia. 
The general results of these investigations are that a decrease in heat 



3 I 4 

production does not play any important part in the maintenance of 
body temperature. 

The action of "high temperatures and humidity upon the respira- 
tory gas exchange has been investigated mostly under artificial. 
conditions in a temperate climate, and the results are not very 
definite, and in some cases contradictory. Briefly all that can be 
said is that there appears to be an ill-defined zone of temperature, 
about 15 0 to 25°C. (59 0 to 77 0 F.) in which the metabolism is at a 
minimum. 

The effects of extreme humid heat upon the respiratory 
metabolism have been studied by Harvey Sutton (1908). When the 
wet bulb temperature of the surroundings rose above a critical point, 
as previously shown by Haldane, so that no evaporation of sweat 
could take place, the body temperature rose, and this caused an 
increased metabolism and corresponding increased production of 
heat, which again further increased the body temperature; a vicious 
circle was thus established, resulting in a gradually increasing rate 
in the rise of body temperature, until dangerous conditions were 
reached. 

Eijkman (1896), working under natural conditions actually in the 
tropics, determined the respiratory exchange of eleven Europeans 
and twelve Malays, and compared the average amount of oxygen 
consumed and carbon dioxide expired with the averages obtained 
by workers in Europe. Using the Zuntz-Geppert method, he 
obtained, as the result of thirty-seven experiments on Europeans and 
forty-eight on Malays, an average of 245*7 c.cs. of oxygen absorbed 
per minute by the Europeans, and 215*5 by the Malays; these he 
compared with figures obtained by the same method in Europe by 
Geppert, Loewy and others, namely, 250*3 c.cs. per minute. In 
both cases the numbers are calculated to a body weight of 
64 kilograms. He concluded from these that there was no evidence 
of any decreased heat production. It is obvious that the number of 
experiments is far too small to draw definite conclusions. 

Observations were carried out, also under natural conditions, by 
Osborne (1912) during the summer in Victoria, Australia, where the 
temperature frequently rises above ioo°F., and he found a decided 
increase with the higher external temperatures, both in the pulmonary 
ventilation and in the carbon dioxide produced. 



315 


The reaction of the metabolism to surrounding conditions has 
been investigated recently by Hill and his colleagues (1914) in 
England, by determining the respiratory exchange from day to day 
under natural conditions, and co-ordinating the changes he observed 
not with the actual air temperature, but with the cooling power of 
the atmosphere as determined by the katathermometers, two 
thermometers with large bulbs filled with alcohol. This instrument 
was invented by Hill to measure the rate of cooling due to 
atmospheric conditions on a body at a temperature in the 
neighbourhood of that of the human body. The two instruments 
employed (dry and wet bulb katathermometers) give the rate of 
cooling, the dry one by convection and radiation, and the wet by 
convection, radiation and evaporation. He found that in experi¬ 
ments done in the open air, the greatest metabolism took place on 
days on which the greatest rate of cooling was observed. He also 
found that in experiments done indoors where the rate of cooling 
was low, the metabolism was very markedly less than outdoors upon 
the same day, provided that the cooling power outdoors was very 
much higher than indoors. In short, the metabolism ran parallel 
with the cooling power of the atmosphere, and not with the actual 
temperatures as observed with the ordinary dry and wet bulb 
thermometers. He attributes the efficacy of the open air treatment 
of pulmonary complaints to the increased metabolism produced by 
the greater degree of cooling outside. 

The present paper contains the results of experiments extending 
over three years, the subjects living in the tropics (19 0 South), with 
the object of investigating the changes in the respiratory metabolism 
under different atmospheric conditions. Observations have been 
carried out in the hot season of the year (November to May) and in 
the cooler season (June to October). In these experiments a 
similar procedure was followed to that of Hill, the cooling power 
of the atmosphere being observed by means of the katathermometers 
as well as the temperatures with the ordinary dry and wet bulb 
thermometers. 

The writer wishes to acknowledge his indebtedness to Mr. J. W. 
Fielding, chief assistant of this Institute, who not only acted as one 
of the subjects, but helped with the various observations recorded. 



3 f 6 


METHODS EMPLOYED 

The subject lay at ease in a deck chair, with a foot rest, and 
remained without movement during the experiment. The usual 
canvas of the chair was removed and replaced by a covering made 
from an open-mesh string hammock, so that the body* should be 
freely exposed on all sides. After lying for about thirty minutes, 
the subject breathed for exactly twenty minutes through a mouth¬ 
piece fitted with inspiratory and expiratory valves into a Zuntz 
portable respiration meter, the nose being closed by means of a 
spring clip. The meter was previously tested by driving through 
it a known volume of air from a large aspirator at the rate of between 
5 and 6 litres per minute, and a correction determined from the 
mean of a large number (ioo) of such tests, and applied to the 
volume of expired air in each experiment. This correction was very 
small, amounting to an addition of 2 c.c. per 100 c.c. measured. 
Certain modifications were employed in collecting the sample of 
expired gases for analysis. A mixing vessel was interposed 
between the mouthpiece and the meter, as recommended by Bohr 
(see Krogh (1916)), consisting of a cubical metal box of about 
2 litres capacity. The sample was drawn out through the usual 
opening attached to the small box at the top of the meter, but the 
proportionate sampling arrangement employed by Zuntz was 
replaced by a gas sampling tube as designed by Huntly (sec 
Haldane (1912)), which allowed of the sample being collected at an 
even rate over the whole period, merely by allowing the mercury with 
which the sampling tube was filled to run out through an exit tube 
of a suitable bore so as to collect about 180 to 200 c.c. in the time. 
The water acidulated with sulphuric acid, used by Zuntz, was thus 
dispensed with, as it was found that this absorbed a small quantity 
of carbon dioxide, especially if the analysis were postponed 
for any time. The sample was analysed by means of the Haldane 
apparatus, at least two analyses of each sample being performed. 

Table I shows a series of pairs of observations, done at different 
times to test the method, each pair consisting of consecutive twenty 
minutes* breathing into the meter. The two observations were 
substantially the same in each case. 



317 


Table I. 


Subject 

Vol. of 
expired air 
Litres per 
20 min. 

Analysis o 
A 

f expired 
ir 

CO a 

eliminated 

c.c. 

per min. 

O a 

consumed 

C.C. 

per min. 

Resp. 

Quot. 

CO.% 

o,% 

w. J. Y. (1) . 

no 

366 

16-50 

. ! 99 . 

254 

0-78 


112 

369 

16-51 

205 

260 

o -79 

(*) . 

108 

349 

1674 

186 

236 

0-79 


-hi 

3*37 

* 6-55 

l 97 j 

250 

°79 

(3) ... ... 

87*5 

372 

l6-28 

161 

214 

0-75 


86*o 

3-80 

l6-C>9 

162 

220 

. °*74 

J. W. F. 

120 

3*6 7 

l6-82 

217 

253 

o-86 


116 

! 

3 ’ 5 8 1 

16*89 

205 

241 

0*85 


Whilst the breathing was going on, the following observations 
were made by the assistant at the beginning, half-way through, and 
at the end of the time, and the averages of the readings so obtained 
taken as the figure for the whole time: — 

1. The readings of the ordinary dry and wet bulb 

thermometers. 

2. The temperature of the expired gases in the meter. 

3. The temperatures, taken • with dry and wet bulb ther¬ 

mometers, of the air between the skin and the clothing 
(called by Hill skin-shirt temperatures). 

4. The times taken for the dry and wet bulb katather- 

mometers to cool from no 0 to ioo°F. and ioo° to 
90°. As a rule, only one determination was made with 
the dry bulb instrument, as the time taken was generally 
too long to allow of a series of observations. 

5. The pulse rate was noted, and the rectal temperature of the 

subject was taken before and after each experiment. The 
pulse rate did not show any great variations, the two 
extremes with W. J. Y. being 64 and 76 per minute, and 
with J. W. F. 60 and 70 per minute. 

The volume of expired gases was calculated dry to 760 mm. 
pressure and o° C., and for the calculation of the oxygen absorbed, 
a correction was made for the difference in volume of inspired and 
expired air as given by Haldane (1912). The percentages of 
oxygen and carbon dioxide in the inspired air were taken as 20 90 



318 


and 0 03 respectively, the figures given with outside air by the 
Haldane gas analysis apparatus employed in the experiments. 

The experiments were carried but on two subjects, W. J. Y., 
38 years at commencement and 45*4 kilograms in body weight, and 
J.W.F., 27 years and 56'2 kilograms body weight, and were 
always done in the mornings about the same time after breakfast. 
Neither subject altered in weight during the whole period by more 
than a ftw tenths of a kilogram. A number of preliminary experi¬ 
ments were performed in order to get the subjects accustomed to 
the mouthpiece. 

In all but a few experiments the same amount of clothing was 
worn, consisting of a light shirt, cotton trousers, socks an<J shoes. 
Most of the experiments out of doors were carried out in the shade 
of a large mango tree, and a few on the verandah of the first storey 
of the Institute. The accompanying tables give the details and 
results of the experiments. 


AIR TEMPERATURE 

The climate in the coastal districts of North Queensland is 
monsoonal, and the rains occur during the hot months of the year 
(November to April), during which time the readings of the wet 
bulb thermometer are very high, and typical moist tropical weather 
is experienced. During the hot season the outside dry bulb 
temperatures lay between 8o° and 91 0 F. (267° and 32‘8° C.), and 
the wet bulb was with a few exceptions always between 75 and 
8o°F. (23*9° and 267° C.), whilst higher wet bulb temperatures 
, (over 8o° F.) were observed indoors. In the cooler season experi¬ 
ments were carried out with much lower air temperatures, the lowest 
dry bulb reading being 62‘8° F and the lowest wet bulb 5 i' 4 ° F. 

• 

THE SKIN-SHIRT TEMPERATURE 

It is a well-known fact that there is always a layer of stagnant 
air between the clothing and the skin which is usually at a higher 
temperature and at a greater degree of saturation with moisture than 
the external air. The more the body perspires and blocks the pores 
of the clothing the greater becomes the degree of saturation of this 



air. In a cool climate the difference in temperature between this air 
immediately surrounding the body and the outer air is much more 
marked. 

For comparison with the experiments of Hill, dry and wet bulb 
temperatures were taken by two thermometers which were prevented 
from touching the skin by being enclosed in a small wire cage. The 
bulb of one was covered with a woven cotton glove and was dipped 
in water before inserting between the skin and the shirt. The 
figures obtained in these experiments under various conditions are 
given in the tables. 

It was observed that when the subjects were out of doors the 
skin-shirt temperatures ran roughly parallel with the air tempera¬ 
tures, and were several degrees above these. In the hot season the 
difference between the dry bulb temperatures of the skin-shirt air 
and the outer air was less marked than between the wet bulb 
temperature. 

Hill has published a number of skin-shirt temperatures which he 
collected in England, and it is noticeable that many of his figures 
taken out of doors in winter with the subject at rest, are as high as 
those registered in North Queensland in a tropical summer (e.g., he 
observed wet bulb temperatures of 88° on several occasions). It 
must be borne in mind that the subjects of Hill’s experiments were 
wearing heavy clothing, whereas the clothing worn here was of the 
lightest description. These figures thus show the influence of the 
clothing on the temperature of this air, and bear out the truth of 
Rubner’s remark that a clothed man always lives in a tropical 
climate as far as his body is concerned. 

The effect of moving air upon the skin-shirt temperature were 
also studied by Hill who showed that these greatly effected this 
temperature; on a windy day he recorded skin-shirt temperatures 
as low as 64° F. wet and 74 0 F. dry, whilst temperatures lower than 
these were also observed by him when the rush of air was made by 
cycling rapidly downhill into the wind. The effect of air movement 
in the experiments done in Townsville was not nearly so marked as 
in those quoted above. 

The figures recorded in the cooler season of the year show much 
lower skin-shirt temperatures. A wet bulb skin-shirt temperature 
as low as 64*8° F. was observed on a very windy, showery day in 



320 


June, the subject at the time feeling so cold that he was shivering. 
Much higher temperatures were observed when the subject was in the 
sun; thus, on one occasion skin-shirt temperatures of 103 6° dry bulb 
and 99*6° wet bulb were observed, and on another 98*7° dry and 
90*7° wet bulb. 


THE KATATHERMOMETERS AND RATE OF COOLING 

It has been recognised that the wet and dry bulb thermometers 
do not give a true indication of the effect of the atmosphere upon 
the human body. Anyone who has lived in a hot moist climate 
knows the immense difference in comfort between moving and still 
air, yet little change may be observed in the actual air temperatures 
under the two conditions. Since the comfort or discomfort of the 
body is determined by the rate at which cooling can take place, and 
not by the actual temperature of the surroundings, Hill has devised 
the katathermometers, which measure the rate of the heat loss, i.e., 
the cooling power of the atmosphere under the existing conditions. 
In the tropics when the outer temperatures are high, the evaporation 
of the sweat plays a much greater part in the cooling mechanism of 
the body than it does in a temperate climate, so that‘the rate of 
cooling of the wet bulb katathermometer is a closer representation 
of the actual conditions than the dry bulb instrument. 

In these experiments the time taken for the dry katathermometer 
to fall from 1 io° to ioo° only was generally recorded, on account of 
the long time taken to cool to 90°: the wet bulb was generally 
carried on to 90°. These times are given in the tables in seconds. 
It has been found convenient for comparison of the figures to express 
the rate of cooling over the whole ten degrees in degrees lost per 
minute; these figures are also given in some of the tables. 

It was observed By Osborne (1916) in Melbourne, that the wet 
bulb katathermometer is extremely sensitive to air currents, and that 
successive readings outdoors, especially in dry weather, may show 
very considerable variations. Katathermometer observations carried 
out here have shown this objection to be justified, especially as 
regards the wet bulb instrument. If a series of readings be taken 
in rapid succession in still air indoors, or outdoors on a still day. 



321 


fairly consistent times may be obtained, and the same is true if a 
steady wind be blowing. On a day, however, with the wind coming 
in gusts great variations may be observed. Experience in 
Townsville has shown how very seldom days occur in which a series 
of observations can be done out of doors without fairly large varia¬ 
tions, and what would be felt as a still tropical day is often not so 
calm when tested by the wet katathermometer. Several observations 
with the wet katathermometer are given in Table II to illustrate 
these points. 

Table II. 


No. 

Air Temperature 

Wet Bulb Katathermometer 


Dry B. 

Wet B. 

110°—IOO° 

ioo°—90° 




seconds 

seconds 


(0 

881 

817 

76 

*52 

Indoors with doors 


89*0 

819 

73 

* 5 * 

closed. 10 minutes 


89*0 

81-9 

77 

*55 

readings. 

« 

633 

52-0 

*7 

20 

Outdoors, cool season 



... 

*5 

20 

breeze blowing. 


647 

5>'8 

>5 

22 

5 minutes readings. 


658 

52-8 

16 

21 


(3) 

8 7 -i 

80*2 

67 

128 

Hot season. Outdoors, 




48 

89 

wind in gusts. 


... 


52 

83 

5 minutes readings. 


... 


76 

*24 


(4) 

74-0 

66*3 

39 

60 

Outdoors, cool season. 


75-2 

66*9 

46 

56 

Breezy day. 


76*2 

67*3 

32 

5 * 

10 minutes reading. 

■(5) 

82-4 

77*2 

56 

104 

Day felt extremely close 


82*0 

765 

49 

74 

and uncomfortable. 


. 81-8 

7 6 ’3 

57 

94 

5 minutes readings. 


82-0 

77*0 

62 

99 



82-0 

77 *° 

61 

* *4 



Nos. i and 2 show that constant readings could be obtained 
indoors, as well as outdoors when a strong steady wind was 
blowing; Nos. 3 and 4 show the effects of gusts of wind, and No. 5 
shows the variations over half an hour on a day that was classed 
to the senses as very close and uncomfortable. 

On account of these variations a series of observations was made 
with the wet katathermometer at intervals during the whole time of 
each experiment, and the average of the series taken as a represen- 


322 


tation of the cooling power during the time. With the dry 
katathermometer the time occupied in cooling, especially in the hot 
season, was much longer; thus, a. sudden gust of wind would not 
produce so noticeable an effect. 

The times recorded for the katathermometers to cool, in most 
cases, are of course much longer than those recorded by Hill in 
England. Indoors, in the hot season, the average time taken for 
the wet bulb katathermometer to cool from iio° to ioo° was 
74 seconds, and for the dry bulb instrument 330 seconds, 
approaching in fact some of the observations recorded by Hill in an 
artificially heated chamber, and similar to conditions found by him 
in weaving sheds in England, where the wet bulb temperature was 
8o°F. and more. 

In the outdoor experiments considerable variations were noted, 
the rate of cooling from 1 io° to ioo° varying from 250 seconds dry 
bulb in the hot season to 42 in the cool season, and 60 seconds wet 
bulb down to 16 seconds in the cool season. 

It is interesting to note that on one or two days in June the time 
of cooling of the katathermometers was quite as low. as some of the 
times recorded by Hill in England in winter (Table V, No. 16, and 
Table VI, No. 12). 


THE RESPIRATORY METABOLISM 

The experiments carried out indoors, Tables IJI and IV, were 
all performed in the same room with doors closed, so that in the 
comparison between the two seasons the only differences in outer 
conditions were those due to atmospheric temperature. Indoors, 
during the hot season experiments, the air temperatures averaged 
87° F. dry bulb and 8o° F. wet bulb, whilst in the cool season the 
averages were much lower, 71 0 to 74°F. dry bulb alid 63° to 67° 
wet bulb. The rate of cooling indoors, as measured by the 
katathermometers, did not vary to any great extent on the different 
days of each season, but a marked difference was observed between 
the average rates of cooling during the hot and cool weather. With 
both subjects the average quantities of oxygen consumed and carbon 
dioxide eliminated were greater in the hot season. Thus, in subject 
W. J.Y., the average oxygen consumed was 248 c.c. in the hot 



Subject W. J. Y. Indoor Experiments. 


323 







3 2 4 


season, and the average carbon dioxide eliminated was 200 c.c., 
whereas in the cooler weather series averages of 204 and 219 for 
the oxygen and 155 and 168 for the carbon dioxide were obtained. 
Taking, for convenience, the minimum absorption of oxygen in each 
series as a basis, the extreme variation observed in the hot season 
series was 23 per cent., and in the two cool seasons series 8 per cent, 
and 16 per cent, of the minimum respectively. It is noticeable 
further, that the total quantity of air breathed in this subject was 
distinctly greater with the higher external temperatures. 

With the other subject, J. W. F. (Table IV), similar observa¬ 
tions were recorded indoors, the average for the oxygen being 256 
and 222 c.c., and for the carbon dioxide eliminated 217 and 183 c.c. 
respectively in the hotter and cooler weather. ' 

In the experiments done out of doors the external conditions 
were more varied. Large variations were observed in the wet and 
dry bulb air temperatures in the hot season, and the cooling power 
of the atmosphere, as measured by the katathermometers, also 
showed considerable differences from day to day. Thus, with the 
air temperatures above 8o° dry and 70° wet bulb, the extreme 
variations in the time that the dry bulb katathermometers took to 
cool from uo° to ioo° were 250 and 94 seconds, and the wet bulb 
instrument 60 to 35 seconds, whilst in the Cooler weather, with lower 
air temperatures, times taken were on more than one occasion as low 
as 45 seconds for the dry bulb and 19 seconds for the wet bulb 
katathermometers. In these outdoor experiments, however, it was 
also noticeable that the metabolism was generally higher with the 
higher outer temperatures. 

The comparison was rendered more difficult in the outdoor 
experiments by the fact that on certain days in the cooler weather 
the subject felt decidedly cold, often sufficiently so to be shivering, 
and on these occasions an increased metabolism was observed 
(Table V, Nos. 14, 15, 16 and Table VI, Nos. 10, 11, 12). An 
example is No. 16 in Table V, done on a windy day in June with 
air temperatures of 64*3° dry bulb and 51 *4° wet bulb, in which the 
subject was so cold as to be shivering, in spite of the fact that he 
was wearing a coat; the rectal temperature fell o*8° during the 
period. On these days it was observed that the temperatures 
between the skin and shirt were much lower. 



Subject J. W. F. Indoor Experiments. 








Subject W. J. Y. Outdoor Experiment*. 


326 




3 2 7 


Another question investigated was whether the cooling' power 
of the atmosphere observed during the hot season, as measured by 
the katathermometer, influenced the metabolism in any way. The 
figures did not show any definite changes in the metabolism with 
rate of cooling. In the cooler weather a high rate of metabolism 
was observed in a few cases on days which had a high rate of 
cooling, but it was noticeable that this occurred only when the 
subject felt cold. 

No such marked changes were observed as those obtained by 
Hill in England, who found in outdoor experiments that the 
greatest rate of metabolism coincided with the greatest rate of 
cooling. 

Observations were made on the effects of increasing the rate of 
cooling by performing experiments, first indoors and then outdoors, 
upon the same day, in the same manner as done by Hill (1912). 

The subject in each case reclined motionless for at least 
40 minutes under each set of conditions before breathing into the 
meter. Tables VII and VIII give the results of a number of such 
experiments, the indoor experiments being carried out in a closed 
room. In the hot season experiments no increase in the metabolism 
was observed out of doors, although the cooling power was 
considerably higher outside. The difference in comfort indoors in 
a closed room and outdoors in the breeze was very marked. Moving 
air was thus without any definite effect when the outer temperatures 
were high (over 8o° dry bulb and 70° wet bulb), showing that the 
increased rate of cooling was insufficient to reduce the body 
temperature during the experiment. With the lower external 
•temperatures observed in the cool season differences were observed. 
The pulmonary ventilation was generally greater out of doors, and 
in certain instances there was a decided increase in carbon dioxide 
eliminated and oxygen consumed. This occurred on windy days 
when the cooling power of the atmosphere was high and when the 
subject felt cold: on these days the rate of cooling was often 
sufficient to reduce the rectal temperature. On the whole, however, 
no marked increase in the oxygen consumed was observed, ^nd it is 
noteworthy that the averages for the eleven observations on W. J. Y. 
showed no increase out of doors excepting that the total volume 
expired was greater. In several experiments the increased ventila- 



Tabli VL 


328 







Subject W. J. V. Effects of Wind. Experiment! done Indoors and Outdoors. 


329 


Remark*. 

In 

Out 

In 

Out 

In 

Out 

In 

Out 

In Cooler season. 

Out Feeling cold ; strong 

In breeze ; Rectal temp. 

Out fell to 98°. 

In 

Out 

In 

Out Very little breeze. 

In 

Out Gusty breeze. 

In 

Out Good breeze; feeling cold. 

In 

Out Very slight wind. 

In 

Out Moderate wind. 

In 

Out Wind in gusts. 

In 

Out Good breeze. 

In 

Out Wind in gusts; uneven 

In cooling. 

Out * 

lN 

Out Very strong wind. 

In 4 days in hot season. 

Out Nos. 1—4. 

All indoor experiments hot season 
ix observations. 

All outdoor experiments same 

period 11 observations. 
In 2 days in cool season, 1918 

Out Nos. 5 and 6. 

In 11 days in cool season, 1919. 

Out Nos. 7—17. 

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33i 


tion may have caused a washing out of carbonic acid from the 
tissues, since on some occasions the respiratory quotient was slightly 
higher out of doors. The difference in the rate of cooling indoors 
and outdoors was on no occasion so great as on those days recorded 
by Hill as ‘bracing/ when he observed a very large increase in the 
metabolism out of doojs. The rate of cooling due to air movement 
must therefore be increased beyond a certain limit in order to 
stimulate the body to an increase in respiratory metabolism. These 
results agree in the main with Wolpert’s observations that moving 
air has very little effect upon the carbon dioxide output between 
78° and 95 0 F., but below this it produces an increase. 

One experiment was carried out under similar conditions to those 
investigated by Harvey Sutton (1908) with high temperatures and 
so high a saturation of moisture as to prevent all cooling through 
evaporation of sweat. This was done in a small chamber heated 
artificially. Before entering the chamber the subject (W. J. Y.) 
breathed in the usual manner for five minutes into the meter, and a 
sample was taken for analysis. After thirty minutes in the hot 
room, a second breathing into the meter was performed, and again 
after seventy minutes. The figures obtained are seen in Table IX. 


Table IX. 


Air Temp. 

Wet-bulb 

Rectal 


Vol. 
expired 
Litres 
per min. 

CO* 

o a 



D.B. 

W.B. 

Kata. 

IIO°-IOO C 

Temp. 

Pulse 


m 

Resp. 

Quot. 


81-6 

695 

63 

99*5 

76 

6*4 

198 

216 

0*91 

Before entering. 

97-8 

93.0 

j 

... 

100-5 

112 

8*6 

294 

327 

0*90 

After 30 mins, in 
hot room. 

100-3 

96*0 

241 

0 

^ • 

0 

168 

97 

3 11 

375 

083 

After 70 mins. 


The subject was naked and was perspiring profusely during the 
experiment, losing 650 grins, in weight, practically all of which 
would be due to sweat. The result was similar to those recorded 
by Sutton, the rectal temperature rose, slowly at first, more rapidly 
later, and the metabolism was increased. It was not observed, 
however, that the respiratory quotient rose towards unity as 







33 2 


described by Sutton. The last breathing was extremely trying, the 
mouthpiece proving somewhat uncomfortable under the existing 
conditions, and was accompanied by a sensation of suffocation, 
which may have affected the breathing in this case. 

It is interesting to note here some observations which were made 
during exercise. The effort consisted in walking on the flat for 
fifteen minutes, at the rate of three miles per hour, and the 
meter was carried on a wooden stand strapped to the shoulders 
like a knapsack, the total weight carried being approximately 
13 kilograms. These experiments are detailed in Table X. The 
temperatures given are the shade temperatures at the time; the 
walking, however, was performed in the sun. The calories, 
corresponding to the oxygen consumed, are calculated from the data 
given by Cathcart (1915). 


Tablf X. 



Shade ' 

Dry B. 

Pcmp. 

Wet B. 

Rectal 

Temp. 

Vol. 
expired 
Litres 
per min. 

COa 

c.c. 

per min. 

! 

i 

°* , 
C.C. 

per min. 

Resp. 

Quot. 

Resp. 

Rate 

Calories 
per min. 


W. j. V. (.) 

83-8 

75 -o 

98-8 

4*9 

167 

I 

220 

076 


1*06 

Rest. 




996 

19*4 

781 

959 

o*81 

19* 

462 

Exercise. 

( 2 ) 

84-0 

73 ’° 

98 '4 

4*9 

*74 

211 

0-82 

124 

1*02 

Rest. 

■ i 



1002 

20-7 

901 

1119 

o*81 

26! 

5*40 

Exercise. 

J. W. F. (3) 

00 

73 1 

99-1 

6-8 

228 

256 

0-89 

* 4 * 

1*26 

Rest. 




998 

174 

698 

836 

0*83 

204 

404 

Exercise. 

( 4 ) 

83-4 

73*9 j 

997 

6*6 

218 

2 54 

o*86 

i6* 

1*24 

Rest. 




100*0 

20-2 

I 

1007 

1260 

0-80 

20 J 

6*05 

Exercise. 


As already pointed out, in the tropics the lo§s of heat from the 
body takes place to a greater extent by evaporation of the 
perspiration than it does in a temperate climate. It is of interest, 
therefore, to calculate the amount of water which would have to be 
evaporated to neutralise the heat produced in these experiments, 
assuming for the time that all heat was lost'in this way. 

The largest production of heat with subject W. J. Y. amounted 
to 5 4 calories per minute; i litre of water evaporated at 98’6° 




333 


requires 582 calories, so that to neutralise this amount of heat would 
require the^ evaporation of about 9 c.c. per minute or 540 c.c. 
per hour. 

With the second subject (J. W. F.) in Experiment No. 4, 
6*5 calories were produced, and the evaporation of 10*4 c.c. of 
water per minute would be required to neutralise this heat, or 
626 c.c. per hour. 

IiT another research (1919) the loss in body weight of the same 
two subjects during walking exercise was measured, the subjects 
being weighed in their clothing, so that the loss in weight 
practically represents the perspiration evaporated. The first 
subject (W. J. Y.), in walking on a hot day (D. B. 87*8°, W. B. 
79*7°) for sixty minutes at about three to three and a half miles per 
hour, lost 740 grins, in weight, and the second subject (J. W. F.), 
walking the same distance in seventy-six minutes, lost 900 grms. 
or about 710 grms. per hour. 

It is thus seen that even with a high moist temperature sufficient 
evaporation can take place to neutralise the heat produced by such 
exercise as that performed, even if this evaporation were the only 
source of heat loss. These experiments, however, caused profuse 
perspiration, and it shows how necessary it is in the tropics to 
supply the body with sufficient water (cf. Hunt (1912)). 


DISCUSSION 

The ventilation of the lungs, carbon dioxide expired and oxygen 
consumed, were greater in the summer than in the cooler weather, 
excepting on a few days when the subjects were cooling sufficiently 
rapidly to feel cold and to produce shivering, on which occasions 
larger metabolism was observed. 

The results agree, therefore, with those found b’y Osborne (1912), 
who found that the pulmonary ventilation and the carbon dioxide 
produced varied directly with the external temperatures, and who 
pointed out also that the rate of respiration is noticeably higher with 
higher external temperatures (over ioo°F.). 

An increased rate of respiration in the tropics has been noticed 
by other observers; $ius Chamberlain (1911), as the result of 
extended observations on six hundred and eight American soldiers 



334 


stationed in the Philippine Islands, found an average respiration 
rate of 19*3 per minute, as against 17 to 18 per minute in temperate 
climates. An increased rate of respiration would in itself cause a 
certain small increase in the carbon dioxide produced. 

With one of the subjects (W. J. Y.) the rectal temperatures in the 
cool weather were in most cases lower than in the summer, which 
would account for a certain difference in the metabolism; with the 
other subject, however, this was not the case, the rectal temperature 
did not show any marked variation in the two seasons, being 
generally about 99 0 in all the experiments. Osborne also states that 
he was unable to observe any rise in rectal temperature whilst at rest 
which would have caused the increase observed in the metabolism. 

A discussion of the various conditions which may affect the basal 
metabolism, i.e., the metabolism with complete muscular rest and in 
the post-absorptive condition, has been published by Benedict 
(1915), in which he sums up the results of the large number of 
experiments which have been carried out in the Nutrition Laboratory 
of the Carnegie Institute, Washington. These experiments have 
been made on a large variety of individuals, and have extended 
over a number of years. He pointed out how numerous the factors 
are which may affect the basal metabolism. Besides those more 
obvious factors directly connected with the mass of the organism, 
such as body weight, body surface, muscular development, etc., 
there are others which, though less obvious, have a marked effect. 
He has noted considerable changes (as much as 30 per cent.) from 
day to day in the oxygen absorption of normal individuals, and 
small changes in the course of twenty-four hours even in fasting 
subjects, which changes could not be attributed to changes in the 
active mass of the body. 

Benedict and Cathcart (1913), experimenting with a professional 
athlete, found that after a prolonged period of severe work the 
metabolism remained high, although gradually decreasing, long 
after the external evidence of muscular activity had ceased. They 
further found that the metabolism of this individual, determined 
from day to day, showed variations which in some cases might be 
traced to conditions existing prior to the experiment; for instance, 
when he chose a rough road on his way to the laboratory the extra 
exertion affected the subsequent determination, and the same was 
true in the winter, when his walk was impeded by slippery roads. 



335 


Benedict concludes, therefore, that besides the active mass of the 
organism, ‘ the stimulus to cellular activity 1 existing at the time the 
measurement of the metabolism is made, is an important factor in 
determining the metabolism observed. 

The metabolism at any time is thus determined not only by the 
conditions which obtain at the time of the actual experiment, but 
also by the conditions which have gone before, the results of 
which on the cellular activity of the organism have not had time to 
disappear. 

The foregoing observations suggest an explanation at any rate 
for part of the increased metabolism found in the summer in these 
experiments. In a tropical summer with a hot moist atmosphere, 
when the least exertion is accompanied by profuse sweating, a very 
slight amount of muscular work will cause an increase in the body 
temperature and a corresponding increase in the metabolic activity. 
The increased activity produced by the exertion of dressing and 
walking to the laboratory would persist, and would not be 
materially reduced by the subsequent rest, before and during the 
actual measurement. In the summer here, it has frequently been 
noted how rapidly the body temperature rises with exercise and falls 
again only very slowly when the exercise has ceased. It was 
frequently observed that the subject was perspiring freely during the 
experiments. In the cooler season the effects of the earlier exertion 
would not be so great, and the greater cooling power of the 
atmosphere would further cause the resting period to modify the 
activity to a greater extent. The body temperature during the hot 
weather seldom changed very much during the actual experiment, 
whilst in the cooler season in the outdoor experiments it often 
showed a decided fall. The amount of actual exertion before the 
experiments began was roughly the same every day, yet the body 
temperature of one subject was usually greater in the hot weather. 

One might thus expect to find the metabolism working at a 
slightly higher level in the very hot weather. It may be noted here 
that very cold weather might possibly produce a similar effect, since 
the increased muscular activity produced by the cold might also 
persist for some time. 

That the metabolism was not wholly influenced by the immediate 
conditions is also seen by the fact that during the winter the rate 
of cooling of the katathermometers inside with doors closed was of 



336 


the same order as that frequently found out of doors in the summer, 
although the oxygen consumed was lower in the former experiment. 

The fact that the level of metabolism is affected by the general 
conditions under which the subject is living, and not only by the 
immediate experimental surroundings, has been entirely overlooked 
in experiments of comparative short duration, in which tropical 
conditions have been produced artificially, and it shows how these 
can never replace observations carried out under natural conditions 
upon subjects living continuously under these conditions. 


SUMMARY 

During the hot season of the year a greater metabolism was 
observed than in the cooler season, excepting on certain days in the 
latter, when the rate of cooling was sufficiently great to cause the 
subject to shiver. This greater metabolism is attributed to the 
effects of the ordinary activities of everyday life which had preceded 
the actual experiment, which in the hot moist weather produce a 
greater increase in body temperature, and consequently in the 
metabolism, than in the cool season. These effects also are reduced 
much more slowly during the hot season, when the cooling power of 
the atmosphere is low, than during the cool weather when the 
cooling power is much greater. The effects of such conditions prior 
to the actual measurement have thus a larger influence on the level 
of metabolism in the hot season. 

With tropical heat the metabolism is at a high level on account 
of the increase in body temperature produced by even slight 
exertion, and which decreases only very gradually after the 
exertion has ceased Cold may also increase the metabolism but by 
producing shivering and so increasing the muscular activity. 


337 


REFERENCES 

Benedict (1915). Journ. Biol. Cbcm. Vol. XX, p. 263. 

Bkkinl and Young (1919). Med. Journ. of Australia. Vol. VI, i, pp. 353, 375 and 395 
[reprinted in this issue]. 

Cathcart (1918). Journ. R.A.M.C. Vol. XXXI, p. 339. 

Chamberlain (19 i i). Philipp. Journ. Sci. B. Vol. VI, p. 427. 

Eijkmann (1896). Pfliigers Archiv. Vol. LXIV, p. 57. ' 

Haldane (1912). Methods of Air Analysis. London. 

Hill, L. (1914). Local Government Board Reports. New Series, No. 100. 

Hunt (1912). Journ. Hyg. Vol. XII, p. 479. 

Johansson (1896). Skand. Arch. Physiol. Vol. VII, p. 123. 

Krogh (1916). The Respiratory Metabolism of Animals and Men. London. 

Loewy (1890). Pfliigers Arch. Vol. XLVI, p. 189. 

Osborne, W. A. (1910). Journ. Physiol. Vol. XLI, p. 353. 

Osborne, W. A. (1916). Proc. Roy. Soc. Vic. New Series. Vol. XXIX. 

Sutton, Harvey (1908). J^urn. Path, and Baci. Vol. XIII, p. 68 . 

Rubner, and Lewaschew (1897). Arch. f. Hygiene. Vol. XXIX, p. 1. 

Wolpert (1902). Arch. f. Hygiene. Vol. XLIII, p. 236. 




339 ■ 


THE HYPOPUS OF CARPOGLTPHUS 
ANONYMUS, Haller 


BY 

HUBERT M. MORRIS, M.Sc. (Manch.). 


(Received for publication 12 December , 1919) 


A quantity of dried figs received from the Port Sanitary 
Authority, Liverpool, were found to be heavily infested with mites 
and small beetles in all stages of development. The mites were 
found to be Carpogl-yphus anonymus (Haller), and the beetles were 
Carpophilus hemipterus (Linn). 

A few days later a single hypopial nymph was seen. 

No other species of mites were present, and as the hypopus shows 
some points of similarity to the other stages of Carpoglyphus 
anonymus, there seems no doubt that it belongs to that species. 
Only one example of the hypopus was met with amongst a great 
number of specimens of the other stages. 

This hypopus is broader in relation to its length, less strongly 
convex dorsally, and much more active than the normal nymphs of 
the species. 

It has been considered worth while to figure and describe this 
hypopus, as Michael (1903) states that it is unknown. 

The hypopial nymph forms a very interesting stage in the life 
of those species of mites in which it occurs. Its occurrence appears 
to be entirely confined to mites belonging to the family 
Tyroglyphidae, and it has not been recognised in all members of 
that family. 

The hypopial nymph occurs apparently between the two normal 
nymphal stages, or after the first nymphal stage where only one 
occurs; the hypopial period being preceded and followed by an 



34 ° 


ecdysis in the usual way. By no means all individuals become 
hypopi, but usually only a relatively small number, and in some 
species, as probably is the case with that which is now described, 
it is of rare occurrence. The hypopi may develop into adults of 
either sex. 



Their occurrence does not seem to be caused by unfavourable 
conditions, but is a provision for facilitating the distribution of the 
species, and to that end they are adapted for being transported by 
insects, larger mites, and other animals, by the development of 
relatively powerful suckers and in some cases large claws, as well 
as by being usually considerably more active in this than in other 
stages of their development. 


The habit of the hypopi clinging to insects, etc., has frequently 
led to their being considered as parasites, but as they do not prey 
upon their host, but merely adhere to it in order to be transported 
from place to place, they are not true parasites, although on 
- occasions when they occur in enormous numbers, as is sometimes 
the case, one may safely infer that they are deleterious to their host. 

The integument of the hypopus differs from that of the adult, 
being of such consistency as to afford them greater protection, 
enabling them to endure greater degrees of heat and drought, such 
as they are liable to meet with while undergoing transportation. 


DESCRIPTION OF THE HYPOPUS OF CARPOOLYPHUS 
ANONYM US, Haller. 

Colour: Translucent white, the legs very slightly tinged with 
pink. 

Form: Broadly ovate, approaching to circular owing to its 
relative width. Dorsally it is rather convex, but ventrally it is 
flattened. 

The cephalothorax projects between the first pair of legs as a 
blunt triangle, which bears two pairs of hairs, and below this is a 
very slightly developed rostrum. 

A pair of short hairs are situated laterally at the posterior end 
of the cephalothorax; two short pairs at the posterior end of the 
body and a pair anterior to the sucker-plate. 

The first pair of epimera are joined to the sternum, the other 
pairs are free. 

The sucker-plate is strongly chitinised, and nearly circular, but 
is truncated posteriorly. The plate appears to be divided in the 
median line in front. At the anterior end of the plate there are a 
pair of suckers of medium size, close together; behind these are 
another rather larger pair also close together, lateral to these 
another pair of smaller suckers, and posterior to them an additional 
pair. 

The legs are approximately equal in length, each bearing a well 
developed caroncle and a slightly developed claw. The legs also 
bear a number of hairs, of which the majority are short and slender. 



3+2 


The tibia of the first leg bears a hair about equal in length to that 
of the entire leg; the tibia of the second leg bears a similar hair, but 
it is only about half the length of the leg; the third and fourth legs 
each bears a similar and still shorter hair; there is also a well 
developed hair on the tarsus, which is longer than that on the tibia, 
particularly in the case of the fourth leg. 

The integument bears a number of striations which on the 
cephalothorax lie roughly transversely, and on the abdomen in a 
more or less longitudinal direction. 

The length of the specimen was 0 24 mm. and its breadth 
o’20 mm. 


REFERENCE 


Michael (1903). British Tyroglypbidae , Vol. II, p. 44. 



343 


XEROPHTHALMIA IN A NATIVE OF 
THE GOLD COAST 

BY 

J. W. S. MACFIE 

(Received for. publication 7 January , 1920) 

In a recent number of the Journal of Tropical Medicine and 
Hygiene , Archibald (1919) described three cases of epithelial xerosis 
of the conjunctiva in natives of the Sudan, a disease which, he 
remarked, apparently had not been previously described as occurring 
in the Tropics. 

Epithelial xerosis of the conjunctiva, or Xerophthalmia, is a not- 
uncommon affection of natives in West Africa, but, as a rule, it 
causes no trouble, and so does not come to the notice of Medical 
Officers. It is for this reason presumably that its occurrence has not 
been previously recorded. Occasionally it happens, however, that 
a patient becomes alarmed about the white patches on his eyes, 
fearing that they may spread and obscure his vision. A case of 
this sort was sent to me a short time ago, and the opportunity was 
taken of attempting to transmit the disease to experimental animals. 


CLINICAL HISTORY OF THE CASE 

The patient was a native of Accra, Gold Coast, 23 years of age, 
a fitter by trade. He stated that six years ago he had an illness 
which he was told was ‘ Bright’s disease/ the chief symptoms being 
cough, blood in his urine, and swelling of his legs. Soon after this 
illness white patches appeared on his eyes, and had persisted ever 
since. The white patches caused neither pain nor discomfort, were 
not spreading, and did not interfere with vision; but it was evident 
that the fear of blindness had become an obsession. The conjunctiva 
was not particularly dry; indeed, the patient complained that the 
secretion from his eyes was sometimes excessive, a condition which 
he attributed to dust and metal filings getting into his eyes when 
at work. There was no night blindness. 



344 


The general health of the patient was good. His diet was that 
customary for an adult native of his class, and was ample in every 
respect. His urine contained a considerable quantity of albumen, 
a circumstance which may have had something to do with the 
condition of his eyes. 

On the conjunctiva of each eye, external to the corneal margin 
and a little below the middle line, there was a triangular dull-white 
j>atch, the superficial layers of which were easily scraped off, 
revealing brown pigmentation underneath. Direct smears made 
from the scrapings from the patches showed the usual cellular 
elements, and large numbers of Gram-positive, diphtheroid bacilli 
resembling Bacillus xerosis . 


CULTURAL CHARACTERS OF THE BACILLUS 

After washing one eye thoroughly with sterilised normal saline 
solution, a little of the white patch on it was scraped off with a 
sterile needle and used for inoculating media. In this way a pure 
culture of a bacillus was obtained which on agar grew slowly, at 
the end of forty-eight hours showing minute colonies, raised, 
rounded, and semi-transparent; on ‘Nasgar* showed a growth 
similar to that on agar; and in broth produced after forty-eight 
hours a slight turbidity and a little fine granular deposit. The 
bacillus was a non-motile, Gram-positive, diphtheroid organism 
which was not acid-fast, and did not form spores. 


BIOCHEMICAL REACTIONS OF THE BACILLUS 

No change was produced in maltose, dextrin, inulin, starch, 
salicin, inosite, glycerol, erythritol, dulcitol, iso-dulcitol, mannitol 
or sorbitol. Acid was produced in glucose, laevulose, lactose^ 
saccharose, galactose; and slight acidity in amygdalin. Gas was 
not liberated in any of the media employed. These tests were made 
in Hiss* serum-water medium, containing I per cent, of the carbo¬ 
hydrate or other substance, inoculated from a culture of the bacillus 
growing on blood serum. 

The biochemical reactions of the diphtheroid bacilli found in the 
eye vary greatly; those of the organism isolated frota this case 



345 


differed from those of the bacillus recovered by Archibald from his 
Case I, the patient from whom a successful inoculation was made on 
to the eye of a rabbit, since the latter produced acid in galacfose, 
rhamnose, maltose, lactose, erythritol, and dulcitol. 

PATHOGENICITY OF THE BACILLUS 

The bacillus was not pathogenic to guinea-pigs when inoculated 
sub-cut aneously. 

Although diphtheroid bacilli are found in great numbers in the 
lesions of xerophthalmia, they are not, as a rule, regarded as the 
cause of the disease. This question, however, has been re-opened 
by Archibald, who succeeded in producing a xerotic area on the 
conjunctiva of a rabbit by applying to an abraded surface ‘an 
emulsion of the viscid xerotic material* obtained from the eye of one 
of his cases. As it was possible that such material might contain, 
besides B. xerosis , other things which did not grow in the 
cultures, this procedure was not followed, but instead an emulsion 
was used of a culture growing well on agar after several sub¬ 
inoculations. 

A little of the emulsion was applied to the eyes of the following 
animals after gently scratching the conjunctiva with a needle—one 
goat, one sheep, three rabbits, two guinea-pigs, two tame rats 
( E. rattus) and two monkeys (Cercopithecus patas). No xerotic 
patches were produced. The observation period was three weeks in 
the goat, sheep, one rabbit, and one guinea-pig, and ten weeks or 
longer in the other animals. 


SUMMARY 

(1) Epithelial xerosis of the conjunctiva is a not-uncommon 
affection of natives in West Africa. 

(2) Attempts to reproduce the disease in animals, by means of 
the bacillus obtained from a case of this disease, were unsuccessful. 


REFERENCE 

Archibald, R. G. (1919-) Epithelial Xerotii of the Conjunctiva in Natives of the Sudan. 
Journ. of Trop. Med. & Hyg., XXII, p. 81. 




347 


AN OBSERVATION ON THE EFFECT OF 
MALARIA IN LEUKAEMIA 

BY 

J. W. S. MACFIE 
(.Received for -publication 7 January , 1920) 

On a previous occasion (1917) it was shown that in a case of 
lymphatic leukaemia a great reduction in the number of leucocytes 
took place when the patient developed malaria. The leucocytes, 
which had numbered as many as 286,000 per c.mm. before the 
attack, were reduced to 59,000; but the ultimate effect was not 
observed, as the patient left Accra for k more healthy district and 
was consequently not seen again. The effect of an attack of malaria 
in leukaemia has again been observed recently, on this occasion in 
a case of the myelogenous type. For this reason, and because the 
examinations were in some respects more complete, a brief record 
may be of interest. 

The patient was a native boy, aged 18 years, suffering from 
myelogenous leukaemia. He complained of nothing but a little pain 
and ‘hardness* on the left side of the abdomen. His spleen was 
greatly enlarged, reaching downwards to a point two inches below 
the umbilicus; and he was anaemic. The superficial lymphatic 
glands were not palpably enlarged. He had not recently suffered 
from fever, and dated back the onset of his illness only six weeks. 
He was first examined by me on the 5th of August, 1919, and was 
subsequently seen weekly until the 9th September; that is, six 
examinations were made at intervals of seven days. 

The treatment given was tartar emetic, to which was added 
quinine when it was found that the patient was infected with malaria. 
The details were as follows: — 

Intravenous injections of tartar emetic, one grain each, on 
August 6th, 9th, nth, 13th, 17th, 19th, 22nd, 25th, 28th, 
31st, September 2nd, 4th, 6th, and 8th—total 14 grains. 
Quinine sulphate, 30 grains daily, by the mouth in solution, 
from August 20th to September 9th. 

The results of the blood examinations are shown in the Table and 
in the Graph. It will be noted that the number of leucocytes per 



348 

Successive examinations or the blood in a case or myelogenous leukaemia. 


Date of 
observation 

Blood Cc 
per c. 

trpusdes 

nun. 

Examination 

for 

Malaria parasites 

Remarks 

White 

Red 

. 

August • 5 th 

■ 326,250 

3,047,000 

Neg. 

Intravenous injections of tartar 
emetic started on Aug. 6th. 

12th 

104,870 

2 > 45 , > 34 0 

P. falciparum : 
not numerous 

Morning temperature on 

Aug. nth, ioi° F.; on 

Aug. 13th, ioo° F. 

„ 19th 

62,900 

3 > 334 > 4 °o 

P. falciparum : 
numerous 

Quinine sulphate, 30 grains 
daily, orally, started on 

Aug. 20th. 

„ 26th 

162,130 

3 » 43 2 > I 5 ° 

Neg. 


September 2nd 

290,940 

3,200,000 

Neg. 


» 9 th 

364,37s 

2,906,250 

Neg. 




















349 


c.mm. was reduced from 326,250 to 62,900 between the 5th of 
August and the 19th, a phenomenon which might have been 
attributed to either the treatment with antimony, which was Started 
on the 6th, or to the intercurrent malaria (. P . falciparum ), which 
developed about the 12th of August. On the 20th of August 
quinine was given for the first time. Subsequently malaria parasites 
were not again found in the blood, and although the antimony 
treatment was continued the number of leucocytes increased steadily, 
so that on the 9th of September they were more numerous than they 
had been at the time of the first examination. 

In myelogenous leukaemia, just as was previously observed in 
lymphatic leukaemia, a notable reduction in the number of leucocytes 
coincides with the appearance in the blood of malaria parasites. 
This reduction is not permanent, and is rapidly effaced by treatment 
with quinine. 

Incidentally this case shows also, (1) that intravenous injections 
of tartar emetic in the doses given have no beneficial action in 
myelogenous leukaemia, and (2) that such injections do not prevent 
the development of an attack of malaria ( P . falciparum ). 


REFERENCE 

Mac m, J. W. S. (1917.) A Note on the Occurrence of Leukaemia in the Native* of West Africa. 
Report of the Accra Laboratory for the year 1916, pp. 39-42. J. and A. Churchill, 
London. 




35 ' 

TROPICAL AUSTRALIA AND ITS 
SETTLEMENT 

BY 

i 

A. BREINL 

AND 

W. J. YOUNG 

{From the Australian Institute of Tropical Medicine , Townsville ) 

Reprinted from Medical Journal of Australia, Vol. VI, i, 

PP- 353 * 375 and 395. 

CONTENTS pagk 

I. Introduction .351 

II. The Climate of Northern Australia .353 

III. Sunlight in the Tropics ... ..363 

IV. Effects of Heat and Humidity on the Animal Organism . 368 

(1) Regulation of body temperature under artificial conditions ... 369 

(a) Changes in the physiology of the white man living under 

TROPICAL CONDITIONS .3 JZ 

( a ) Body temperature in the tropics.374 

( b ) Rats of respiration -.376 

(c) Blood pressure.. .377 

( d ) Blood conditions of Europeans in the tropics ...' ... 377 

(?) Metabolism in the tropics. . .380 

(/) Effect of tropical climate on the nervous system . 389 

( g ) Clothing in the tropics . 391 

(3) Summary of physiological changes .394 

V. White Settlement in Tropical Australia .397 

(1) General conditions in Tropical Australia and statistics ... 397 

(a) Housing .403 

(3) R*sum* .407 

I. INTRODUCTION 

Tropical Australia and its settlement by a British race has 
formed the theme of numerous public utterances and of a good deal 
of writing in both the scientific and lay press. Sometimes these 
utterances have been dictated by utilitarian reasons alone, some- 



times they have only formed part of a general political propaganda; 
many have been influenced by personal prejudice, and were put 
forth by writers whose experience of life in Northern Australia did 
not qualify them to express an opinion on this subject. 

On many occasions, when the ultimate fate of Northern 
Australia has formed the subject of discussion at various scientific 
meetings, the speakers have applied indiscriminately to Northern 
Australia the outcome of experiences gained by themselves or by 
others in various parts of the tropics, without realising that Northern 
Australia occupies a different position, on account of the absence 
of a settled native population and its peculiar climatic conditions. 

The following pages are the outcome of several years* investiga¬ 
tion in and consideration of conditions obtaining in Northern 
Australia and their influence upon a settled white race. A r6sum6 
is included of previous investigations by other workers into the 
practicability of a white settlement of other parts of the tropics and 
the influence of tropical life upon a population of European descent. 
An attempt is made to sift facts and to review the results of previous 
investigations, published in numerous scattered journals, most of 
which, with the exception of the experiments carried out in the 
Philippine Islands, not only have been done in a haphazard way, 
but sweeping conclusions have been based upon a few unsystematic 
observations. 

The problem of the settlement of tropical Australia by a white 
race is a very difficult and complicated one, since two factors enter 
into the question. In the first place, climatic conditions and their 
effect upon the white man are of paramount importance, since racial 
degeneration, brought about by climatic conditions alone, would 
decide the problem. At the same time the economic side of the 
question plays an important part and cannot be neglected, and a 
discussion on this aspect has been included. 

Opportunity is taken here to thank Mr. H. A. Hunt, the 
Commonwealth Meteorologist, for putting at our disposal the 
climatic data, and also Mr. G. H. Knibbs, C.M.G., the Common¬ 
wealth Statistician, for -help and suggestion in connexion with the 
statistical data. 



353 


II. THE CUMATE OF NORTHERN AUSTRALIA 

The Australian living in the southern parts of his continent 
generally possesses only a very uncertain knowledge of the climate 
of the northern parts. He assumes that the whole of the north above 
the tropic has a uniformly hot climate, that heat and mugginess 
persist and that a feeling of personal Gomfort is only a rare 
sensation. This popular conception arises from the general idea of 
'tropics 1 ; most people in their early youth associate with 'tropics* 
a land of impenetrable jungle, heat, swamp and fever, an impression 
gained by reading books of adventure and travel. 

The scientific study of climates within the tropics shows, 
however, that there exists a great diversity of climatic conditions, 
regulated by a number of factors, such as the nearness to the ocean, 
elevation above sea level, proximity to high mountain ranges which 
control rainfall, prevailing wind and, above all, the amount and 
monthly distribution of the rainfall. 

To describe climate from the point of view of its effects upon a 
human race is impossible with our present means. It is only to a 
certain extent that climate, as we feel it, finds a graphic expression 
in those meteorological observations usually recorded, which measure 
heat and humidity only, namely, the readings of the dry and wet 
bulb thermometers. From these readings is further calculated the 
relative humidity, that is, the extent to which the atmosphere is 
laden with moisture. The degree of personal comfort or discomfort 
experienced is indicated to some extent by these readings, but it 
is certain that many other factors play a large part in determining 
individual sensations. 

Numerous attempts, summarized as far as 1908 by Hann, have 
been made to construct a ' discomfort scale,* but no satisfactory 
solution has yet been attained. Conspicuous amongst these is that 
of Cleveland Abb 4 , who suggested a ‘curve of comfort,* based 
upon three factors, namely, air temperature, relative humidity and 
wind velocity. By plotting temperatures against humidities for a 
certain velocity of wind and estimating personal comfort or 
discomfort under these conditions, he obtained charts which corre¬ 
lated his personal sensation with the above factors. 

Wet bulb temperatures alone were first used as a guide to comfort 



35 + 


by Harrington, who termed them ‘sensible temperatures* and 
mapped out wet bulb isotherms for the United States of America 
for the month of July. The importance of wet bulb readings has 
been further emphasised by Haldane (1905), who demonstrated 
experimentally that the regulation of body temperature above a 
certain wet bulb reading became disorganised and human energy 
was paralysed. He therefore suggested wet bulb temperatures as a 
standard by which to regulate conditions in factories, mines and 
workshops. 

Griffith Taylor (1916) has suggested a tentative scale of 
discomfort, applying only to warmer regions, where humidity and 
temperature are the chief factors, a scale depending on wet bulb 
readings. He divided ‘climate* into the following grades: — 

8*3°-i27° C. (45°-55° F.) wet bulb—Most comfortable. 

I27°-I5 , 5° C. (55°-6o° F.) wet bulb—Very rarely uncomfortable. 

* I5*5 0 -i8*3£ C. (6o°-65° F.) wet bulb—Sometimes uncomfortable. 

i8’3°-2ri° C. (65°-7 o° F.) wet bulb—Often uncomfortable. 

2i*i°-237 0 C. (70°-75° F.) wet bulb—Usually uncomfortable. 

Over 237 0 C. (over 75 0 F.) wet bulb—Continuously uncomfortable. 

A rough-and-ready indication of the degree of discomfort, used 
by Osborne (1916), is the amount of clothing required to enable him 
to lie in an open-meshwork hammock. His experiences have led 
him to set a wet bulb of about 22*75° C. (73 0 F.) as an empirical 
standard above which truly tropical conditions arise. 

Many authors, however, contend that wet bulb readings alone 
are not an unerring indication of the subjective sensation caused by 
climate. L. Hill (1914, 1916), for example, pointed out: ‘It is no 
use to trust to the ordinary thermometer, either wet or dry, because 
it does not show heat loss. It only shows its own temperature, the 
average temperature of the furniture, surroundings and walls; it 
does not show the heat loss of the body. Whilst the thermometer 
is a static instrument indicating average temperature, the human 
body is a dynamic structure, continually producing and losing heat, 
while its temperature remains sensibly constant.* He deprecated the 
use of relative humidity as a criterion of comfort, and suggested the 
katathermometer, a new apparatus, which would register the rate 
of heat loss and thus indicate more closely the effect of temperature 
and atmospheric conditions upon the human body. This instrument 



355 


consists of two thermometers, each having a large bulb filled with 
alcohol; one of these is kept dry—the dry bulb katathermometer; 
the other is kept moist by means of a wet cotton glove—the wet bulb 
katathermometer. Both are heated to 43'3°C. (no°F.) and the' 
time which each takes to cool from 43*3° to 32 2 0 C. (1 io° to 90°F.) 
is noted. The dry bulb katathermometer thus measures the rate of 
cooling by convection and radiation, the wet bulb katathermometer 
that by convection, radiation and evaporation and thus measures 
not the actual air temperature but the rate of cooling of a moist 
body, due to the effects of the atmospheric condition at an actual 
temperature approximating to that of the human body. Practical 
experience has shown, however, that the wet bulb katathermometer 
is too sensitive to air currents. Osbome (1916) has pointed out that 
a series of readings taken in rapid succession yields fairly uniform 
results only in still air indoors, or outdoors if a steady breeze be 
blowing. On the other hand, when the wind comes in gusts, 
consecutive readings show great variations. Observations in 
Townsville with the katathermometers have confirmed Osborne’s 
experience, and have proved that calm weather, as far as the 
katathermometer readings are concerned, is a rare exception, and 
days classed as still to personal sensation may not be so to the 
katathermometer. 

Griffith Taylor (1916) attempted to overcome in a novel manner 
the difficulty of judging a climate, by introducing a graphic 
representation, termed a climograph, and plotting the monthly 
means of relative humidity against monthly means of wet 
bulb readings. He compiled as a standard a composite 
climograph, by using average figures for towns situated in 
regions where human energy is at its best, selecting for this 
purpose five towns in the southern and seven towns in the northern 
hemisphere, where the average monthly wet bulb readings ranged 
between 27 0 and i66°C. (37 0 and 62° F.). This climograph, 
according to Taylor, represents ideal conditions for the white race, 
and he terms it the ‘white race climograph.’ On the same principle 
he constructed a number of climographs for different parts of the 
world and differentiated in this way four extreme climatic types— 
hot and damp, hot and dry, cold and damp and cold and dry— 
and compared the climographs of Australian towns with these types 



356 

(see fig. i). The shape of the climograph also indicates the seasonal 
distribution of rainfall. 

It is an ingenuous and striking method of comparing in a general 
way the climate of any given locality with a type, but does not 
convey any further information than the mean readings of dry and 
wet bulb thermometers, and it is significant that Taylor himself, 
when discussing the question of discomfort in relation to climate, 
uses the wet bulb readings as guide. 



Note.—T he shaded figure is the composite white race climograph based on twelve typical cities. 


Fig. i. 


Bruce (1916) has proposed the use of dew point as an indicator 
of the effects of atmospheric conditions upon the human body, 
and comes to the conclusion that the dew point most desirable 
for human activity is i6’6° (62° F.), that is to say, that air saturated 
with moisture at i6’6° (62° F.) is neither muggy nor chilly. If the 
dew point, however, rises over 2i‘i°C. (70°F.) the conditions 
become exceedingly trying. 



357 


This principle has been applied to the average temperatures 
recorded in Townsville, and the results have indicated that the 
dew point at 9 p.m. is, as a rule, higher than at 9 a.m. or 3 p.m., 
although the conditions in the evenings are, on the whole, 
undoubtedly less trying than those prevalent during the day. 
Furthermore, this principle does not take into account the effects of 
other conditions, such as wind and sunshine. 

It would appear, therefore, that neither humidity nor dew point 
give any more information in regard to personal comfort than the 
readings of dry and wet bulb thermometers. 

Hunt, at Osborne’s suggestion, constructed wet bulb isotherms 
for Australia—the only part of the world for which such complete 
isotherms have been published—attempting thus to represent 
graphically climatic conditions as far as personal comfort is 
concerned. As pointed out previously, wet bulb temperatures alone 
in this respect are only of limited value, since experience has shown 
that, although high wet bulb temperature, approaching the limit of 
safety, may be a guide to discomfort, yet lower ones, without the 
accompanying dry bulb readings, are indefinite. From our personal 
experience, wet bulb readings above a certain limit invariably denote 
discomfort, yet the dry bulb temperature even then modifies the 
degree of discomfort felt. For example, a wet bulb reading of 
26*6° C. (8o°F.) is never pleasant, but the degree of discomfort 
becomes exaggerated in direct proportion to the dry bulb reading. 
For these reasons it has been thought advantageous to make use 
simultaneously of the mean dry and wet bulb readings for the 
comparison of the various parts of Northern Australia. In the 
accompanying graphs (fig. 2) are plated the average monthly dry 
and wet bulb temperatures, taken at 9 a.m. over a period of five 
years, and the points are connected by lines for distinctness only, 
the io°C. (50°F.) line being indicated in order to make a 
comparison easy. Since the temperatures are so dependent on the 
rainfall, average rainfalls obtained from Hunt’s publications have 
been plotted in these charts. The relative positions of the graphs 
of the various towns are arranged diagrammatically as near as 
possible in accordance with their geographical position (compare 
map, fig. 3). Thus the coastal towns are represented on the outside 
of the diagrams and the inland towns in the interior. The graphs 



358 



Fig. 2. 












359 


commence with the month of April, as* in this month a distinct 
change of season sets in; beginning from then, the average tempera¬ 
ture falls (rapidly at first and more gradually afterwards) and begins 
to rise again on or about July. In this way both fall and rise are 
more conveniently displayed for comparison than if the graphs were 
commenced with January in the usual way. In order to make the 
comparison more complete, the monthly averages for the maximum 
and minimum temperatures are displayed in a similar manner on a 
second series of charts (fig. 4). 



Fig. 3. 


The graphs make it clear that the climate of the coastal towns 
differs essentially from that of the inland towns. The former towns 
have, on the whole, higher wet and lower dry bulb readings than 
the latter. With the coastal towns the average readings increase 
gradually with decreasing latitude and the contrast between the cool 
and hot season becomes less and less pronounced. The charts of 
Rockhampton and Thursday Island, the two extreme towns of 
Queensland, situated within the tropics, illustrate this contention. 
In Rockhampton (latitude 23 0 24") the coolest month (July) has 
average dry and wet bulb readings of 16 3° C. (61*4° F.) and 
13*3° C. (56° F.) respectively, and the hottest month (January) has 
readings of 27 0 C. (8o*6° F.) and 227 0 C. (73 0 F.) respectively. The 
average dry bulb temperature is therefore io‘7°C. (19*2° F.) higher, 







360 























361 


and the average wet bulb reading is 9'4°C. (i7°F.) higher in 
January than in July. In Thursday Island, however (latitude 
io° 34'), the corresponding averages are 25 8° C. (78'3 0 F.) and 
23 2° C. (73*8° F.) for July respectively and 28'i°C. (82-7° F.) and 
25'9°C. (787° F.) for January, showing much smaller differences, 
namely, 2'3° and 27°C. (44 0 and 4-9° F.) respectively. In other 
coastal towns, such as Darwin^ Wyndham and Broome, situated on 
the northern coast of Australia, the seasonal variations are more 
pronounced when compared with Thursday Island. During the hot 
season the average readings there are higher than in towns on the 
eastern coast in the same latitude. Broome and Innisfail, for 
example, both situated between 17 0 and 18° South, show this 
difference in a marked degree (see charts), but the geographical 
position alone explains this difference, since Innisfail lies within the 
region of the trade winds, and, in addition, possesses a larger 
average rainfall. 

For comparison only, graphs representing the climates of Daru, 
Port Moresby and Samarai, all in Papua, have been added (fig. 5). 
These graphs do not differ essentially from those of Thursday 
Island. 

In most of the inland towns of Northern Australia weather 
conditions are greatly influenced by altitude, and changes with 
latitude are consequently less pronounced. 

The graphs representing the average maximum and minimum 
temperatures exhibit the same seasonal variations, which in the 
coastal towns become less marked with decreasing latitude and show 
smaller ranges, of temperature. In the inland towns the average 
maximum readings in general are much higher and the average 
minimum readings much lower than in towns on the sea coast. 

A comprehensive account of the distribution of rainfall through¬ 
out Northern Australia has been published by Hunt in a series of 
monographs issued by the Meteorological Bureau. The rainfall in 
North Queensland shows a seasonal distribution corresponding to 
the monsoonal type of climate, the heaviest falls taking place during 
the hot months (December to March), and only occasional showers 
occur during the cooler months. 

There is a ‘wet belt’ on the north-eastern coast, the centre of 
* which lies about Harvey Creek and which extends northward to a 



362 


point beyond Port Douglas and southward as far as Halifax, where 
a very high general rainfall is registered (see Table I). 


Table I.— Rainfall 


Cooktown ... 
Port Douglas 
Cairns 

Harvey Creek 

Innisfail 

Cardwell ... 

Halifax 

Ingham 

Townsville 


181*9 cm. (71*6 inches) 
210*6 cm. (82*9 inches) 
229*9 cm. (90*5 inches) 
426*0 cm. (167*7 inches) 
384*0 cm. (151*2 inches) 
218*6 cm. (86*i inches) 
226*6 cm. (89*2 inches) 
204*5 cm * (8°'5 inches) 
125*2 cm. (49*3 inches 



Fic. 5. 


The inland towns, on the whole, are much drier than the coastal 
towns and show a similar seasonal distribution of rainfall. 























363 


In the coastal towns of other parts of Northern Australia 
conditions similar to those of North Queensland prevail, with the 
one difference that Broome and Wyndham show a comparatively 
small average rainfall, namely, 58 2 cm. and 71*4 cm. (22 9 inches 
and 28* 1 inches) respectively. In New Guinea, Port Moresby and 
Daru, both situated within the monsoonal belt, possess in the same 
way a dry and wet season, whereas Samarai has its rainfall of 
296*4 cm. (116*7 inches) more evenly distributed over the whole of 
the year. 


III. SUNLIGHT IN THE TROPICS 

The main difference between a temperate and a tropical climate 
lies in the greater intensity of the sun's rays in the tropics. This 
greater intensity is solely due to the less oblique path of the rays 
striking the earth, whereby they have thus passed through a smaller 
layer of atmosphere. As a result, a less degree of absorption and 
scattering has taken place, and the chemical and physical activities 
of these rays are therefore greater the nearer to the equator. This 
greater activity of the sun's rays manifests itself in everyday life. 
The newcomer to the tropics soon becomes aware that coloured 
materials, such as curtains, carpets and clothing fade quickly, and 
written matter in ink, when exposed to the sun, gradually gets 
fainter, and, after a time, almost disappears. Similarly, a number 
of chemical preparations decompose, rubber rapidly perishes and 
certain qualities of glass are altered. The frosting of glass flasks, 
microscopic slides and high-power lenses are examples only too well 
known to laboratory workers in the tropics. 

The bactericidal action of sunlight has been known for a long 
time and has been investigated again recently by Clemesha (1912) 
in India, who exposed cultures of faecal bacteria to the sun. In 
other experiments he added large quantities of faeces to water 
contained in a tank with a large surface area (0*6 hectares, or 1 acre) 
expQsed to the sunlight and examined the liquid from day to day 
for bacterial content. Furthermore, he studied the bacteriological 
flora of waters of natural lakes during the monsoonal and dry 
seasons. All these inquiries led him to conclude that the sun has a 
very powerful action in destroying faecal organisms in water. 



364 


Such changes as the above, together with numerous others of a 
similar nature, led to the question of the nature of the rays producing 
these results. Are they brought about by those rays of shorter wave 
length in the violet and ultra-violet portions of the solar spectrum 
(chemical rays), or are they to be attributed to rays of longer wave 
length, situated at the other end of the spectrum, the red and infra¬ 
red rays (heat rays) ? The ultra-violet rays are known to increase 
chemical activity, and many chemical reactions and decompositions 
may be brought about by exposure to these rays. An example is 
the decomposition of oxalic acid into carbon monoxide, carbon 
dioxide and water when exposed to the sun in the presence of a 
uranium salt. This decomposition is almost entirely due to those 
rays of the solar spectrum in the ultra-violet extending from 550/1 
to 291/1 (Freer (1912)). Observers in the Philippines attempted to 
make use of this decomposition of oxalic acid under standardised 
conditions, in order to compare the intensity of the ultra-violet rays 
of the sun in different parts of the world. The results, however, 
showed that the amount of decomposition in the Philippines and 
other parts of the tropics was inconstant and did not bear any 
definite relationship to latitude. The figures obtained by th$ same 
method in a temperate zone were sometimes as high, and even 
higher, than those found in the tropics. 

The extensive observations of the Philippine workers have led 
them to the conclusion expressed by Aron (1911): ‘That the 
spectrum of the sun’s rays does not extend much, if any, further into 
the ultra-violet in Manila than in northern climates.* These 
observers do not agree with Woodruff (1905) and others, who 
attribute the effects of tropical sunlight on the human organism to 
the influence of ultra-violet rays only. Aron (1911) believes that 
‘ the rays of the tropical sun, having greater wave length than those 
in the red and ultra-red end of the spectrum, play the most important 
role in producing the untoward effects generally attributed to tropical 
sunlight *; in other words, he attributes such effects to the heat rays 
alone. 

Gibbs (1912), working in conjunction with Freer and Aron, 
expresses a somewhat similar opinion, and, if altitude and local 
meteoroligical conditions are taken into consideration, he does not 
believe that ‘when the normal intensities are compared, the light of 



the tropics is different from the sunlight of any other regions/ 
Effects upon life in the tropics, usually attributed to sunlight, are, 
in his opinion, due to 4 other meteorological modifications, which go 
to make up climate, namely, duration of sunshine, clouds, rainfall, 
winds and humidity, all of which affect the air temperature; the last 
is probably the most important fact and depends to a large extent 
upon the duration of sunshine/ 

In Manila and Baguio, both in the Philippines, Aron (1911) and 
Gibbs (1912) carried out experiments on animals in order to study 
the effects of exposure to the sun’s rays under varying conditions. 
Different experimental animals, such as rabbits and monkeys, were 
exposed to the midday sun for varying periods. These animals, 
when shielded from draughts, died after an exposure of from thirty- 
four minutes to about one and a half hours, and showed the post¬ 
mortem appearances characteristic of heat stroke. Black rabbits, as 
a rule, succumbed more quickly than light-coloured animals, and it 
was noticed that the sub-cutaneous temperatures, taken by means 
of a thermocouple, in the case of black rabbits, rose quicker and 
higher than in that of the lighter-coloured ones. 

Aron, furthermore, carried out exposure experiments on dogs 
which had been tracheotomised and thus had part of their effective 
heat-regulating mechanism put out of action.* It is well known that 
dogs do not possess sweat glands on the body, but keep their body 
temperature from rising by means of an increased rate of respiration 
and an increased evaporation from their respiratory tract. These 
animals died after about an hour with typical symptoms of heat 
stroke and corresponding post-mortem appearances. In a similar 
manner the body temperature of tracheotomised rabbits rose on 
exposure to the sun more rapidly than that of normal rabbits. 

When monkeys were exposed to an artificial draught during the 
experiment, or were protected by shade from the direct sun rays, 
no injurious effects were noticed. These experiments, according to 
Aron (1911), show that ‘when placed under the fan animals lost the 
excessive heat which reached them by radiation from the sun. Rays 
including the ultra-violet were nevertheless present, and were 

• Considering the relatively small surface of mouth, nose and throat above the tracheo¬ 
tomy wound, compared with the breathing surface of the lungs, it is evident that the operation 
had put out of action only an extremely small portion of the respiratory surface, and cannot 
altogether account for the quicker death of these animals. 



absorbed by the body in the same manner and degree as by that of 
the control monkeys.’ 

These experiments have recently been repeated by Shaklee (1917) 
and further amplified, to ascertain whether it is possible for monkeys 
to become accustomed to the sun (acclimatised) by a gradually 
increasing daily exposure. His results differed from those of Aron, 
as several of his monkeys, even in the beginning of the experiment, 
lived for hours in the direct sunlight without being protected in any 
way. Summarising his results, he states that experimental monkeys, 
exposed to the sun in Manila, may die from heat stroke after varying 
periods, depending to a less extent upon the sun’s rays than on other 
local conditions, such as proximity of a large, hot surface (ground 
or roof), high relative humidity of the atmosphere and low wind 
velocity. Monkeys, however, may become temporarily acclimatised 
by gradually increasing the time of exposure to the sun. This, in 
his opinion, increases the sensitiveness of the nervous mechanism 
which regulates body temperature by an increased rate of perspira¬ 
tion. The fact that administration of atropine, which impairs the 
function of the sweat glands, causes the death even of an acclima¬ 
tised monkey on exposure, is evidence in support of this conception. 

Animal experiments were also carried out by Schmidt, who, 
working in a temperate climate, exposed rabbits to the sun and 
found a rise in the anal temperature which was more pronounced in 
a black than in a white rabbit. In the same way Schilling (1909) 
observed a rise in the skin temperature of rabbits exposed to the sun, 
and several other authors have made similar observations. It must 
be kept in mind, however, that the temperature of rabbits varies 
considerably even under normal conditions, and the struggling 
alone, when the animal is handled, may cause a considerable rise. 

It is not a new observation that animals, even in a temperate 
zone, may succumb to exposure to the sun, as the following example 
illustrates. In a laboratory in the north of England, in which 
monkeys were kept in a glass house, on more than one occasion 
during the summer months several animals died under the symptoms 
of heat stroke, showing an ante-mortem rectal temperature of 43*3° C. 
(iio°F.), which still rose after death. The painting of the glass 
roof with a white wash reduced the inside temperature of the animal 
house and prevented death. 



3 67 


A consideration of the foregoing observations makes it evident 
that exposure to the direct sun’s rays caused such an increase in 
body temperature that the animals finally succumbed to hyper¬ 
pyrexia. In every case animals of dark colour died more quickly 
than those with light fur, on account of the greater absorption of 
heat. 

. The dark skin of most of the aboriginal races in the tropics, from 
the above point of view, would appear to be a disadvantage, and the 
explanation that dark skin affords protection against the effects of 
the siin merely by insulating the body against the deep penetration 
of harmful rays must be modified. 

Attempts to elucidate the rdle played by the pigment in a 
protective sense are not lacking in the literature. Eijkman (1895), 
in Batavia, covered the bulbs of two thermometers with pieces of 
white and coloured human skin and placed the thermometers in the 
sun. He noticed that the brown skin caused a higher rise of the 
mercury (50*1° C.) than the white skin (47*5° C.). 

Aron (1911) and Gibbs (1912), in the Philippines, experimenting 
with live skin, exposed white and coloured persons to the sun and 
recorded the skin temperatures of various parts of the body by 
means of a thermocouple. Their results were somewhat inconstant; 
the skin temperature invariably rose after a time to from three to 
four degrees above normal. Whilst Gibbs found temperatures 
distinctly higher for the dark skin, Aron noted that the white skin 
was always hotter than the brown, and that after prolonged 
exposure the temperature of the brown skin showed a more distinct 
fall. 4 It may be said/ as Freer (1912) remarks in summarising the 
results of both investigators, 4 that as regards rise in temperature on 
exposure to the sun, the white and brown skin (Filippino) are about 
equal, with a slight factor in favour of tho white, but that in regard 
to the very dark skinned negro the temperature on exposure reaches 
a decidedly higher point than it does with either of the others/ In 
its physiological action, on the other hand, the dark skin is superior 
to the white skin. It absorbs a greater quantity of heat rays, warms 
up more quickly and reaches the point where perspiration commences 
earlier and the evaporation of the sweat causes heat loss and 
consequently affects the cooling of the body. 

A consideration of the foregoing experiments and observations 



368 


upon the effects of exposure to the sun suggests that any ill-effects 
are due not to light but to heat. These experiments, however, only 
take into consideration those effects known as sun stroke or heat 
stroke, and it is, moreover, an almost impossible task in such 
experiments to study the physiological action of the other rays with 
the entire exclusion of heat rays. 

The effect of sunlight on living organisms has formed the subject 
of many publications, and many opinions have been put forward 
which are not founded upon facts but are only wild speculations. 
Woodruff (1905), for example, wrote an extensive monograph on the 
effects of tropical light on the white man, and his conclusions have 
been widely quoted in the literature. He contended that the ultra¬ 
violet rays of the tropical sun are inimical to white settlement, but, 
unfortunately, he based many of his arguments on false premises, 
and many of his statements were merely expressions of personal 
opinion. As a glaring example, the following sentence may be 
quoted : * The southern hemisphere, except the tip of Patagonia, is 
north of 45 0 and therefore unfit for blondes (he assumes that a 
blonde race cannot live nearer to the equator than 50°), and even in 
New Zealand and Australia the native white families are already 
dying out or kept alive by constant new importation from home.* 
He assumes, further, that in New Zealand there is ample evidence 
of the physical decay of the white population. In view of such 
statements, entirely without foundation, it is difficult to consider any 
of his conclusions seriously. 


IV. EFFECTS OF HEAT AND HUMIDITY ON THE ANIMAL 

ORGANISM 

In all warm-blooded animals the temperature of the body under 
normal physiological conditions only varies slightly. This tempera¬ 
ture is maintained by two processes—heat production and heat loss. 
The former is brought about by chemical processes, the combustion 
of food material inside the body, the latter by physical means, 
namely, radiation and convection and evaporation of sweat. In the 
clothed human being the heat production plays a much less impor¬ 
tant part in regulation of temperature than the heat loss by physical 



3^9 

means, and it is evident that the extent to which these physical 
means operate must depend on the surroundings. Whereas in a 
cold climate heat loss takes place mainly by radiation and convection 
from the body, in a hot climate, where the air temperature often 
approaches or even exceeds that of the body, the heat loss through 
radiation can only be very small or nil, and heat is mainly lost by 
evaporation of sweat from the skin. 

It was of great interest to inquire into the physiological reaction 
of the animal body to increased outer temperature, and a great deal 
of experimental work has been done in this direction. Attention 
has been paid to the body temperature and the metabolism in 
conjunction with the calorific value of the food, but a great deal of 
this work has been carried out in temperate climates under artificial 
conditions, and comparatively few observations have been made in 
the tropics. 

(i) Regulation of Body Temperature under Artificial Conditions 

Blagden and Fordyce (1775) noticed that their mouth tempera¬ 
tures did not rise above normal when remaining for fifteen minutes 
in a dry room heated to a temperature of II5°C. to I26'7°C., 
although the heat was sufficient to cook a beef steak. Exposure for 
the same length of time, however, in a moist room heated to 54 4° C. 
caused the body temperature to rise to 37‘8°C. In recent times 
Haldane (1905) carried out his classical experiments on the effect 
of humid heat. His observations were made on human beings, both 
at rest and at work, in an artificially heated chamber, as well as 
underground in deep mines, in still air or exposed to artificial 
draughts. He concluded that the temperature of the human 
organism exposed to humid heat begins to rise after a period 
varying according to the individual arid the conditions and then 
continues to do so. The rise in body temperature corresponds with 
the reading of the wet bulb temperature and is practically indepen¬ 
dent of the dry bulb temperature. During rest in still air, wet bulb 
temperature of about 3i i°C. (88°F.) could be borne without any 
abnormal rise in rectal temperature, but when the thermometer 
rose above that temperature the rectal temperature immediately 
commenced to rise, which rise was accompanied by an increase in 



37 ° 


the pulse rate, profuse sweating and dyspnoea, until finally 
exhaustion set in. During muscular exercise in still air this increase 
in body temperature commenced at a much lower wet bulb (27°C., 
or 8o‘6°F.,) and in moving air much higher wet bulb temperatures 
were required to produce the same effect. 

These experiments were continued and amplified in Haldane’s 
laboratory by Harvey Sutton (1908), who studied the changes in 
the gaseous metabolism of the body when exposed to humid heat. 
He observed a large increase in the metabolism, running parallel 
to the rise in body temperature; in fact, analogous conditions to 
those existent in fever. He concluded that ‘all experiments point 
to the fact that once the balance of the mechanism of heat regulation 
in the human body has been definitely upset by high external 
temperatures, combined with almost total abolition of heat loss in 
evaporation, a vicious circle is established. The internal tempera¬ 
ture rises and as a result the oxidation processes-and therefore 

the production of heat also increases—so that the body temperature 
rises still further, and so on. Once the ball has been set rolling, 
nothing seems to check its progress, and it slowly but surely gathers 
speed. The rectal temperature not only continues to rise, while the 
external temperature still remains constant, but the rise gradually 
increases in rapidity as if momentum were being gathered.’ 

- Conditions similar to those under which the experiments of 
Haldane and Harvey Sutton were conducted, may, according to 
Pembrey (1913), give rise to heat stroke. He analysed the history 
of a large number of cases of heat stroke which had occurred amongst 
British troops in India, and came to the conclusion that heat stroke 
is brought about by a failure of the mechanism which regulates the 
body temperature, after prolonged exposure to excessive moist heat. 
In an excessively hot and moist atmosphere the cooling effect of the 
evaporating sweat is insufficient to prevent the body temperatures 
from rising, and a prolonged exposure to these unfavourable circum¬ 
stances leads to a gradual rise in body temperature and ultimately 
to heat stroke. 

Observations on the influence of wind and draughts on the body 
bring additional evidence to show that, with high external tempera¬ 
ture, the loss of heat due to evaporation of sweat is the most 
important factor in regulating the body temperature. As far back 



37i 


as 1883 Herman suggested that the ill-effect* due to bad ventilation 
and crowded rooms were caused by heat stagnation. This view 
was confirmed experimentally by Heyman, Paul and Ercklentz 
(1905), and later still by Hill, Flack, Rowlands and Walker (1913). 
Hill and his colleagues confined individuals in a small experimental 
chamber until the carbonic acid content rose from 1 per cent, to 
1*5 per cent, in one instance and from 3 per cent, to 4 per cent, in 
another. The subjects experienced discomfort at‘ a wet bulb 
temperature of 277 0 to 28*4° C. (82° to 85° F.) and this discomfort 
was considerably allayed when the air in the chamber was moved 
by a fan. No relief, however, was experienced when fresh air was 
breathed through mouthpiece and tube passing through the wall of 
the chamber. Subjects outside the chamber who breathed the 
chamber air suffered no discomfort, thus proving that the discomfort 
was caused not by the increased carbon dioxide of the air in the 
chamber, but by the heat stagnation, following the failure of 
evaporation of sweat. 

The part played by the evaporating perspiration is illustrated in 
Zuntz and Tendlau's observations upon the body temperature of a 
man whose skin was devoid of sweat glands. The exposure to the 
summer sun of a temperate climate caused his temperature to rise 
to 39° C. (102*2° F.) and a slight amount of manual work produced 
the same effect. He was, however, able to find a substitute for the 
missing function by frequently soaking his shirt in water. 

Such a condition as described above can be artificially produced 
by immersion in a hot-water bath, when the evaporation from the 
skin is prevented and the effects of perspiring are eliminated. Hill 
and Flack (1909) studied the pulse rate, blood pressure, body 
temperature, breathing volume and alveolar tensions of man in a 
hot bath. They found that immersion up to the neck in hot water 
(40*5° to 43’3°C., or 105° to iio°F.) for fifteen to thirty minutes 
caused an increase in the body temperature of 39’ i° to 40*3° C. 
(102*5° to 104*6° F.). The pulse rate and respiration rate and 
volume rose and the blood pressure was lowered. In the alveolar 
air a notable fall in carbonic acid tension and a corresponding rise 
in oxygen tension were observed, caused by the washing-out effect 
of the increased breathing. 

The foregoing observations make it clear that the main difference 



37 2 


between temperature regulation in the tropics and in a temperate 
climate lies in the greater activity of the sweat glands, and the 
evaporation of the increased amount of sweat plays a more important 
part in the cooling of the body. It is a common experience in the 
tropics that the quantity of water secreted in the urine is greatly 
diminished during hot weather, in spite of the larger intake of 
fluid; the balance of the water leaving the body by way of the skin 
and the lungs is considerably larger. 

The establishment of an equilibrium between the water secreted 
by the kidneys and that secreted by the sweat glands is probably 
of importance in the process of acclimatisation. Inhabitants of the 
tropics know only too well the discomfort caused by the frequent 
necessity of micturition during a sudden cold spell, even at times 
when the temperature is well above that regarded as comfortable 
and even warm in a temperate climate. Moreover, the feeling of 
subjective discomfort on such occasions is very marked, and this 
sensitiveness to cold has given rise to the popular belief that the 
blood of dwellers in the tropics is thinner than that of a person living 
in a temperate climate. 


(2) Changes in the Physiology of the White Man Living under 
Tropical Conditions 

The tradition that the white man cannot flourish in tropical 
climes has given rise to a great deal of controversy, and the 
possibility of an acclimatisation has constantly occupied the 
attention of authorities interested in the development of the tropics. 
It has been propounded by several observers with experience of life 
in tropical countries that, apart from diseases peculiar to the tropics, 
climate per se comprising heat, humidity and light make it 
impossible for a white race to settle and thrive. Attempts have 
therefore been made to study the physiology of the white man living 
in the tropics, in order to ascertain whether changes have actually 
taken place, and for comparison the investigations have been 
extended to native races. The results so far are by no means 
complete, and a decided answer is still to seek. 

Little reflection is needed to appreciate the difficulties which 
stand in the way of a practical solution of the question. Tropical 



373 


diseases have up to recent times been very prevalent in most parts 
and have been a large factor in preventing settlement by a white 
race. The pioneer work of Manson and Ross on the r 61 e of insect 
carriers in such diseases as filariasis and malaria has laid down new 
lines for the investigation of other tropical complaints. The 
practical application of the results has led to increased public health 
activities, and has thus created a new era in the permanent settle¬ 
ment of the tropics by inhabitants of European descent. Moreover, 
the tropics, with only few exceptions, such as the South American 
Republics, have only recently been settled; a second and third 
generation is rare, and it is impossible to obtain and select subjects 
in sufficient numbers for carrying out observations which would yield 
results which could not be attacked on the plea of insufficient time 
of observation. The conditions of life in general there, for example, 
housing, ventilation and food, are so different that the difficulty of 
obtaining figures suitable for a comparison of physiological functions 
is greatly increased and comparative standards in many respects are 
lacking. 

A critical review of the literature on acclimatisation brings to 
light the fact that nearly all observations have been made on 
Europeans who have resided in the tropics for a comparatively short 
time. The majority were men chosen for service in the tropics on 
account of their physical fitness, and few of them could boast of an 
uninterrupted residence in the tropics for more than a few years at 
most. 

As an example, in the work of the American observers in the 
Philippines, the figures relating to Europeans were obtained with 
male subjects only, and, furthermore, cannot be accepted, excepting 
on the supposition that changes due to climatic influences become 
established within a comparatively short time. The same criticism 
may be applied to experiments on metabolism carried out by 
Eijkman in Java, whose subjects had resided in the tropics for from 
four and a half to fifteen years, and out of nineteen Europeans only 
one could show a continuous residence for fifteen years. Results 
such as those of Rattray, Wick, Schilling and many others are the 
outcome of observations on one or a few subjects only, collected 
during a flying visit to the tropics. 

Investigations on aboriginal races, although interesting from a 



374 


comparative point of view, do not help to solve the problem, on 
account of the different mode of living, customs and personal frabits. 

(a) Body Temperature in the Tropics. The fact that the 
external temperature in the tropics is considerably higher than 
in a temperate climate suggested a faised body temperature. On 
account of the ease with which the body temperature can be taken, 
it is not surprising that the literature contains records of numerous 
observations, many collected during a sea voyage between Europe 
and the tropics, others on Europeans and native races living in the 
tropics. 

John Davy, in 1839, published a number of observations made 
upon the mouth temperature of seven healthy men during a voyage 
from England to Ceylon, and he concluded that the temperature of 
a European increased whilst passing from a temperate to a warm 
climate, and that, in addition, the body temperature of residents in 
the tropics is slightly raised above normal. He amplified his work 
during a residence of three years in Barbados, and stated that his 
own temperature was higher by o - 5° C. (o'9°F.) when compared 
with his body temperature in England. Many others, including 
Reynaud and Blossville, Rattray, Brown-Sequard, Jousset, Maurel 
and others (referred to by Pembrey, 1898) confirmed this slight rise 
(less than o'55°C., or i°F.), which was more pronounced during 
the first few weeks of residence in the torrid zone, that is, during 
the time of acclimatisation. Crombie (see Pembrey, 1898) recorded 
the results of 1,288 observations upon his own mouth temperature in 
Bengal, and found it slightly higher (namely, o'23° C., or o - 7°F.) 
than in England. Neuhauss (1893), who, during a voyage round 
the world, observed his own rectal temperatures, found the following 
differences (see Table II). 

Tablk If. 


Temperature of Air 

6 a.m. 

12 noon 

1 

| 10 p.m. 

6 p.m. 

1 

Remarks 

Temperate Zone 







Minimum Maximum 







11-5° C. 19-1° c. 

36*6° C. 

3 6 q° C. 

367° c. 

37 '° C. 

Temperate 

Mean 

(5*7° F.) (56-5° F.) 

(97'9° F.) 

(98-5° F.) 

! (98*2° F.) : (98-8° F.) 

in rectum 

>of 20 


55 

55 

56 

62 

Pulse 

J days 

Tropical Zone 


1 

1 

1 




Minimum Maximum 







28*8° C. 26-6° C. 

36-9° C. 

37*3° C. 

3r«°c. 

37-3° C. 

Temperate 

^ Mean 

(75'°° F. ) (79-9° F.) 

(98-5° F.) 

(99*2° F.) 

' (98*8° F.) 1 

(99-*° F.1 

in rectum 

>of 20 


60 

68 

64 

72 

Pulse 

J days 



375 


On the other hand, many observers have denied that there is 
any marked difference in the body temperature; amongst others were 
Thomly (1878) and Fumell (1878), and more recently Wick (1910), 
who could not observe any alteration in his body temperature during 
the passage from a temperate zone to the tropics. 

Chamberlain (1911), as the result of 3,000 mouth temperatures, 
taken at quarterly intervals upon 600 healthy American soldiers 
living in the Philippines, arrived at a similar conclusion. According 
to him, the temperature of American soldiers doing duty in the 
Philippines showed no appreciable variation for season or for 
complexion type; it averaged 37 0 C. (98*7°F.), and this average 
differed little, if at all, from the mean normal temperature for white 
men living in the United States. In Plehn’s (1898) opinion there 
is a slight rise in temperature during acclimatisation, but after 
continued residence this rise disappears. 

The body temperature of dark-skinned native races in the tropics 
does not differ appreciably from that of Europeans. A. Jousset 
(1884) found that the average temperature of Hindoos, Chinese 
and negroes was 37*8° C. (ioo° F.), whereas that of Europeans 
under similar conditions was 37‘87°C. (ioo*i°F.). Eijkman 
(1895) in Batavia compared the axillary temperature of Malays and 
Europeans, and found that the Malays had a slightly lower tempera¬ 
ture (36*85° C., or 98*3° F.) when compared with Europeans (37° C., 
or 99° F.}. 

Young (1915) in a recent paper on this subject recorded a number 
of observations taken during the hot season in North Queensland. 
He pointed out that it is fallacious to speak of a normal mean body 
temperature, as that of a healthy person, even at rest in a cool 
climate, varies considerably during the twenty-four hours,* and 
considered it more rational to employ as standard the range of body 
temperature which has been observed in healthy individuals. He 
concluded that: — 


• Benedict and Carpenter (Carnegie Institution monographs, No. 126, 1910), writing on 
this subject, state that * the idea of constancy in body temperature has become so generally 
accepted that it is commonly believed that the body remains at the temperature of 98*6° F. 
or 37 0 C. without material change, other than that produced by disease. This impression 
is so firmly fixed that one finds to-day on all clinical thermometers a special mark opposite 
this temperature, to indicate what is supposed to be the normal temperature. It is somewhat 
difficult to place the exact history of this mark being selected, and it may be looked upon as 
more or less of a thermometric fetish, which has been worshipped by long-continued usage/ 



1. The temperature of the mouth of Europeans living in the tropics is often 
considerably lower than that of the rectum and that this difference is generally 
much more marked after exercise, just as in temperate climates. The temperature 
of the mouth is thus not reliable as a measure of the body temperature. 

2. During complete rest the rectal temperature did not show any marked 
variations from the limits of temperature observed in Europe. 

3. A considerable rise in the rectal temperature was produced by slight 
muscular work and this was usually maintained for some time after the work had 
ceased. 

These observations make it clear that the discrepancies in the 
records quoted above are in all probability partly due to the mode 
in which the temperatures were taken and partly to individual and 
daily variations. 

At rest the body temperature in the tropics does not vary from 
that in a temperate climate, but with exercise the temperature rises 
more quickly and subsequently decreases more slowly than in a 
temperate zone. A good example of this is quoted by Young in 
respect to one of his subjects, who was in the habit of taking exercise 
in the early morning; during a period of eighteen days his average 
rectal temperature at 8 a.m. was 37*9° C. (100*3° F.). On three 
occasions on which the exercise was omitted the rectal temperature 
at the same time was only 37*5° (99*6° F.). 

(b) Rate of Respiration. A comparison of the respiration rate 
of tropical inhabitants with that found in Europe has yielded 
conflicting evidence. According to Rattray (1870) the total volume 
of air expired was about 7 per cent, less than in a temperate climate, 
and he attributed this to a decreased rate of respiration, since spiro¬ 
meter measurements showed that the lung capacity was increased. 
These observations, made on a small number of subjects, were not 
confirmed by Jousset and by Plehn, who, on the contrary, found an 
increase in the respiration rate. Chamberlain (1911) made observa¬ 
tions on 608 American soldiers in the Philippines. He found the 
average to be 19*3, which is much higher than the figure usually 
accepted as normal for Europeans in Europe, namely, 17 to 18. It 
may be mentioned here that Osborne (1910) carried out observations 
during the summer months in Victoria (Australia), and found that 
with a high outer temperature (37*7° C., or ioo°F.) the pulmonary 
ventilation was increased. 

In consideration of the fact that the human body loses a certain 
amount of water in the expired air and consequently heat due to 



37 7 


the evaporation of the water, it might be expected that the body 
would make use of this means for cooling purposes and an increased 
respiration would be the outcome. 

(Y) Blood Pressure. Estimations of the blood pressure in the 
tropics have been made by a number of observers, with contradictory 
results. Musgrave and Sisson (1910) in Manila examined 97 
Americans, 10 sisters of charity and 40 Filipinos. They graded the 
former two groups according to their length of tropical residence and 
found a decided decrease in the blood pressure, corresponding to 
the length of residence in a hot climate, and suggested that the lower 
blood pressure might be due to a lowered peripheral resistance, 
brought about either by a decrease in the vasometer tone or by 
splanchnic influences. Chamberlain (1911) took the blood pressure 
of 992 American soldiers in the Philippines and made 5,368 observa¬ 
tions, and concluded that ‘the average blood pressure of 115 to 
118 millimetres found in these large bodies of men differed little, 
if any, from the accepted standard among males of the same age 
in a temperate zone when the 5-inch armlet is used/ 

Breinl and Priestley (1914) estimated the blood pressure of North 
Queensland school children, and concluded from 573 observations 
that the climate as such has no marked influence on the blood 
pressure in children. 

Theoretically a permanent change in the blood pressure in the 
tropics should not be anticipated, as a sound heart and sound 
arteries are adaptable to any change. Only temporary variations 
are observed, such as a rise in blood pressure on exertion or a 
lowering due to venous hyperaemia of the skin, similar to that 
produced by a hot bath, but both would only be evanescent and not 
peculiar to tropical life. 

(d) Blood Conditions of Europeans in the Tropics. The 
existence of tropical anaemia has for a long time been regarded as an 
established fact, and most of the early text-books on tropical 
medicine speak of an anaemia due to climatic influences only. There 
is no doubt that the greater proportion of the cases in the time 
before microscopical examination became general were in all proba¬ 
bility caused by malaria and hookworm (anchylostomiasis). The 
skin of the healthy European inhabiting a tropical climate appears 
to the newcomer pale and sallow, the degree of the sallowness 



378 


depending on many factors, especially complexion and skin texture. 
The unprotected skin of individuals with red or fair hair is 
invariably more sallow and appears more anaemic for the same 
reason that it has, on account of the lack of pigment, a lesser 
defensive power against the action of the sun’s rays. The paleness 
of the skin has naturally given rise to the conception that there exists 
a tropical anaemia and that ‘ thinness and poorness ’ of blood is a 
natural sequence of prolonged residence in the tropics. It is not 
unexpected that the advent of accurate microscopical methods has 
led to an investigation of blood conditions of Europeans in the 
tropics, and the results of these investigations have been rather 
surprising. 

Marestang (1889), van der Scheer (1890) and Eijkman (1891) 
examined the blood of Europeans living in the Dutch East Indies 
without finding any marked differences when compared with figures 
obtained for Europe. Glogner (1892) in Sumatra, and Plehn 
(1892) in the Cameroons, carried out similar investigations with 
identical results. These observations, however, are not entirely 
conclusive, since they were based on a comparatively small number 
of estimations. 

Work of a similar nature was carried out by the Americans in 
the Philippines. Wickline (1908) examined the blood of about 70 
American soldiers, after a varying period of residence in the 
Philippines. The first examination was made three months after 
their arrival; it was repeated after the lapse of three months and 
twice again after an interval of approximately eight months. He 
discovered a gradual decrease in the amount of haemoglobin from 
9'7 to 8’3 and a regular rise in the number of erythrocytes from 
4'9 to 5’3 millions. Chamberlain (1911), following up Wickline’s 
work, performed 1,718 red cell counts and 1,433 haemoglobin 
estimations of 702 soldiers, and concluded that ‘ the red cell counts, 
averaging 5 2 millions, rarely falling below 4*5, do not differ from 
the normal at present recognised for healthy young men in a 
temperate zone.’ The haemoglobin figures are perhaps a little low, 
but not sufficient to indicate a definite anaemia. 

Cuthbert (1911) collected similar observations from 21 healthy 
soldiers of European descent on the West Coast of Africa with a 
similar result. Wickline’s and Chamberlain’s figures were obtained 



379 


from healthy subjects especially chosen for tropical service, and after 
a comparatively short residence in the tropics (two years), and are 
open to the objections already mentioned. 

Breinl and Priestley (1914) made observations on apparently 
healthy school children in North Queensland of ages ranging between 
seven and fifteen years, most of whom were bom and had lived all 
their life in North Queensland. Five hundred and eight blbod 
counts were performed on 305 boys and 269 girls, and the haemo¬ 
globin content of the blood was estimated by the Fleischl-Miescher 
method. 

They concluded from their investigations that there is no striking 
difference in the number of erythrocytes and the colour index in 
North Queensland children when compared with averages obtained 
in children of a temperature climate. The number of white blood 
corpuscles (leucocytes), according to Chamberlain (1911), also to 
Sreinl and Priestley (1914), is slightly higher when compared with 
that generally accepted as the standard for a temperate climate. 

Differential counts, which estimate the percentage of the various 
types of leucocytes, showed, according to these observers, a distinct 
alteration. The proportion of the polymorphonuclear neutrophile 
leucocytes appeared to be decreased to 56* 1 per cent. (Chamberlain 
and Vedder (1911)) and 56*8 per cent. (Breinl and Priestley); 
65 per cent, to 70 per cent, is generally accepted as average. The 
percentage of the eosinophile leucocytes was distinctly increased. 

Chamberlain and Vedder (1911) studied the blood conditions of 
American soldiers and natives in the Philippines, and came to the 
conclusion that the Arneth blood picture of the soldiers showed a 
slight ‘shift to the left/ that of the Filipinos a much more marked 
shift, which means that the nuclear condition of the polymorpho¬ 
nuclear neutrophile leucocytes in respect to the numbers of nuclear 
fragments was altered when compared with those of healthy 
Europeans living in a temperate climate; the number of cells with a 
smaller number of fragments (one to two) was considerably increased. 
Breinl and Priestley (1914) carried out analogous observations on 
North Queensland school children, and found that similar conditions 
prevailed and that the blood picture resembled closely that of the 
Filipinos. The authors concluded that the ‘shift to the left* was 
due to the effect of a tropical climate upon the white race living in 
the tropics. 



380 


For comparison a number of Arneth counts on apparently 
healthy native children in the Northern Territory were performed, 
and their ‘Arneth index* corresponded closely to that of North 
Queensland school children. 

The significance of the altered Ameth index is uncertain and, 
according to Breinl and Priestley (1914), the presence of a large 
number of neutrophile leucocytes belonging to class one (possessing 
a single nucleus) ‘ indicates a greater activity of the bone marrow, 
but does not necessarily imply that the. resistance of the organism is 
lowered * (to disease) ‘ in any way.* 

A few observations on the specific gravity of blood were made by 
Eijkman (1896) in Java, and by Breinl and Priestley (unpublished) 
in Townsville. The results did not show any marked deviation 
from the average specific gravity of the blood considered normal for 
a temperate climate, and, when considered in conjunction with the 
previously mentioned blood work, proves that there is in general no 
evidence of an anaemia in the tropics due to climatic influences only. 
The popular belief of the ‘ thinness of the blood * after prolonged 
residence in the tropics is, apart from anaemia due to disease, from 
a scientific point of view only a myth. 

(e) Metabolism in the Tropics. In a cold climate, as has been 
stated previously, the temperature equilibrium of man is maintained 
mainly by regulation of the heat loss through radiation, convection 
and evaporation (termed by Rubner physical heat regulation), whilst 
changes in the heat production (Rubner*s chemical heat regulation) 
take place only to a small extent. In fact, Rubner has said that a 
man feels comfortably warm only when the chemical regulation of 
his heat balance is completely eliminated. In a cool climate 
chemical heat regulation can only be spoken of in the sense that the 
stimulating effect of cold on the skin causes reflex innervation of the 
muscles, resulting in their movement or increased tone, which brings 
about an increase in the production of heat, i.e., the increased meta¬ 
bolism is really due to the muscular action and not to cold per se. 

The question naturally arose whether the chemical heat regulation 
plays any part in maintaining this equilibrium in the tropics where 
the external temperature is high; in other words, whether the body 
responds to these different conditions by a reduced, combustion of 
food material. This problem has been attacked in two ways; first, 



3*1 

by a comparison of the calorific value of the food actually katabolised 
by Europeans living in the tropics, with the average value found in 
temperate climates, and, secondly, by a comparison of the total 
metabolism in both parts of the world, as determined by the 
respiratory gas exchange, that is, the oxygen absorbed and the 
carbon dioxide eliminated, the experiments being carried out other¬ 
wise under exactly the same conditions. 

The most important work- in the first direction is that of Eijkman 
O893), who investigated the food of seven Europeans living in the 
Dutch Indies. His experiments, lasting for several days, were 
carried out as follows : —At each meal each person under observation 
took a second portion of each dish, similar in quantity, as judged 
by the eye, to that which he consumed, and these duplicate meals 
were collected for each day and the protein, fat, carbohydrate, etc., 
in them estimated. At the same time the faeces were collected and 
analysed to determine the quantity of food actually absorbed. 

Allowing for the loss in the faeces, he obtained the following 
figures for the average constituents of the food actually utilised : — 

Protein . 88*2 grins. 

Fat. 79*1 grms. 

Carbohydrate . ... 256*4 grms. 

Ash ... 17*5 grms. 

Alcohol ... 28*5 grms. 

From the results he calculated that for a man weighing from 65 to 
70 kilograms the heat production from the food amounted to 2,400 
to 2,500 calories, a figure which does not differ from that given by 
Rubner for men doing a similar class of work in Euorpe, namely, 
2,445 calpries. The excretion of nitrogen in the urine and faeces 
showed that, on the whole, his subjects maintained nitrogen 
equilibrium during the experiment. 

Eijkman concluded from these experiments that there was no 
evidence that chemical heat regulation played any significant part 
in the thermo-equilibrium of inhabitants of the tropics. 

A different conclusion was arrived at by Ranke (1900), who 
published the results of observations upon himself extending over 
a few days, comparing the calorific value of his food in Europe 
and during a short residence in Brazil. In Europe the heat value 
of his daily food in two experiments was 3,527 and 3,514 calories, 














respectively. In Brazil, however, the calorific value of his food 
was much less, being only 2,812, 1,920 and 1,948 calories, 
respectively, in three series of observations, 

Ranke’s experiments have been criticized adversely by Glogner 
(1909), who conducted similar determinations upon himself at 
different times during his residence in Sumatra and Padang, and 
again after his return to Europe. He stated that loss of appetite 
is a common experience during the first period of residence in the 
tropics, and attributes the lower values obtained by Ranke to the 
fact that the latter had only been a few months in the tropics when 
the experiments were carried out. 

Glogner’s experiments, conducted after a six years’ residence, 
and lasting for fourteen days, showed the average energy value of 
his daily food to be 2,118 calories, or 28*8 calories per kilogram of 
body weight per day. A similar experiment was performed in 
Europe fifteen years later—five years after his departure from the 
tropics—and then the daily average was found to be 2,038 calories, 
or 25 5 calories per kilogram of body weight. It is doubtful, 
however, whether any value can be placed on these observations, 
since the first series was made when the subject was 32 years old 
and weighed 71*1 kilograms, and the series for comparison when 
the subject was 47 years old and weighed 80*3 kilograms. 

Experiments were carried out also upon themselves by Schilling 
and Jaff6 (1909) during a visit to West Africa, and the results were 
compared with observations made in Berlin, both before and after 
their journey. There were no appreciable differences between the 
heat value of their food in both parts of the world, the average 
being 2,863 and 2,953 calories per day for Jaff6, and 2,953 and 
3,021 calories per day for Schilling, in Europe and Africa 
respectively. In both places nitrogen equilibrium was approxi¬ 
mately maintained on about 17 grammes of nitrogen per day. 

In a similar way the nutritive value of the food of native races 
in the tropics has been estimated for comparison with that of white 
people. Eijkman (1893) examined the food of five Malays in a 
similar manner to that employed in his experiments quoted above, 
and found the following average for the food actually katabolised, 
after allowing for the material lost in the faeces: 



3«3 


Protein 

Fat. 

Carbohydrate 
Ash. 


59*9 

25*0 grms. 
462*0 grms, 
13*2 grms. 


The heat value of the food was about 2,349 calories for an 
average body weight of 50 kilograms, i.e., a figure slightly higher 
per kilogram of body weight than that given by. Rubner for men 
in Europe. The amount of unabsorbed matter in the faeces was 
much greater in the Malays than in the Europeans, probably on 
account of the large preponderance of vegetable food in their diet. 

An extensive inquiry has been made by McCay (1912) in India 
into the food of various native races. For Bengalis he found that 
the calorific value of the food was higher (3,196 calories) and the 
nitrogen lower than that of Europeans. A study of the diet and 
nutrition of the Filipinos was made by Aron (1909), and the 
average composition of the daily allowance in the Manila native 
prisons was given as: 


Protein 

Fat. 

Carbohydrate 

Calories 


55*9 g*™- 
27*0 grms. 
510*0 grms. 
2,646 


An estimate of the calorific value of the food eaten by a Filipino 
living under normal conditions corresponded to from 2,500 to 2,600 
calories, and when allowances wqre made for the smaller body 
weight of Filipinos (50 to 55 kilos), this value corresponded to the 
requirements of a working-man in Europe performing moderately 
hard labour. 

Recently, Campbell (1917) examined the food of several native 
students (Chinese, Tamil, Malay and Brahmin), and obtained 
values ranging from 1,502 to 2,492 calories per day. 

The results of these investigations into the food value in the 
tropics make it clear that in general the requirements in calories, 
either of white men or coloured natives, does not differ appreciably 
from European standards. The experiments quoted above indicate 
that the amount of heat produced by the combustion of food is not 
any less in the tropics than in a temperate climate, and that nature 
does not have recourse to a reduced metabolism to regulate body 
temperature. 




384 


The energy involved in the total process of oxidation (heat 
production) taking place in the animal body may be measured either 
by the actual heat value of the food digested (calorific value of the 
food), or again by the quantity of oxygen used up and carbon 
dioxide and water produced (the respiratory exchange), which 
measures the end products of this oxidation. 

If metabolism would play any appreciable part in controlling the 
body temperature in the tropics, it would find its expression in a 
reduced oxygen absorption and carbon dioxide production. 

Experiments upon the influence of high external temperature and 
humidity upon the respiratory exchange have been carried out from 
this point of view under artificial conditions upon mein and animals, 
and on man in the tropics. 

The general -results of these experiments show that with cold¬ 
blooded animals the oxygen absorbed and carbon dioxide eliminated 
are directly proportional to the external temperature; in other words, 
there is no heat regulation, and the body temperature is that of the 
surroundings. With warm-blooded animals, on the other hand, the 
organism regulates its heat to a more or less extent. With the 
monotreme, Ornithorhynchus , the mechanism is almost entirely 
chemical; thus, with increasing external temperature the metabolism 
decreases until a temperature of 32 0 C. (89*6° F.) is reached, the 
body temperature remaining throughout almost constant. Above 
this the animal’s temperature rises, thus indicating that the 
regulatory mechanism is failing, until finally, with increasing outer 
temperatures, the animal behaves like a cold-blooded animal. 

In the rabbit the same is observed, but to a less degree, because 
the mechanism is complicated by loss of heat by physical means. 

In man the elimination of carbon dioxide is not affected by 
rising external temperature in a constant manner. In some experi¬ 
ments, carried out under artificial conditions, it has been noticed 
that with rising temperature the carbon dioxide sinks in quantity 
at a temperature of about I5°C. (59°F.), reaches its minimum 
between 20° C. (68° F.) and 25°C. (77 0 F.), and then rises slightly. 
In other experiments, however, no such change has been observed. 
Generally speaking, all that can be said is that there appears to be 
an ill-defined zone of temperature in which the metabolism is at a 
minimum. When the external temperature and humidity are so 



385 


high that the physical regulation fails, and a rise in body 
temperature results, this rise is accompanied by an increased 
metabolism, still further increasing with continued rise of body 
temperature, until dangerous conditions are reached, as already 
stated. 

A study of the gaseous metabolism under actual tropical 
conditions was made by Eijkman in Java. By means of the 
Zuntz-Geppert method he determined the quantity of oxygen 
consumed and of carbon dioxide eliminated during rest by eleven 
Europeans and twelve Malays. He obtained the following averages 
as the result of forty-eight experiments on Malays and thirty-seven 
experiments on Europeans (see Table III). The numbers are 
calculated to a body weight of 64 kilos. 


Table IIT. 



Oxygen absorbed 
per minute 

Respiratory 

quotient 

Europeans in Java . 

*457 

07QI 

Malays in Java . 

251*5 

0*880 

Europeans in Europe . 

2 5°*3 

°775 


Eijkman interpreted these results as confirmatory of his other 
experiments already quoted, namely, that the heat production by 
the body in the tropics is not less than in a temperate climate, and 
that therefore change in heat production plays an insignificant part 
in the regulation of temperature. He attributes the higher respira¬ 
tory quotient of the Malays to the larger proportion of carbohydrates 
in their diet. 

In reviewing these researches on the metabolism in the tropics 
it is necessary, before drawing any conclusions, to consider the 
limitations of metabolic experiments in general. Contradictory 
results may be accounted for in some measure by different food, 
habits and customs in various countries in which the experiments 
have been carried out, and by individual variations. Moreover, 
up to the present no definite standard has been acknowledged. 

A large number of observations upon metabolism during complete 
rest, which is termed basal metabolism, have been carried out 
during several years in the Nutrition Laboratory of the Carnegie 



386 


Institute in the United States by Benedict and his co-workers. 
Subjects included men and women of various ages, athletes, 
Vegetarians, etc. The general conclusions drawn from these 
experiments have been embodied in a paper by Benedict (1915) upon 
the factors affecting basal metabolism, in which he pointed out that 
a much larger number of factors enter into this question than has 
been hitherto recognised. Although body weight plays an 
important part, there is no direct relationship between the body 
weight and the total heat production. Similarly with body surface, 
the general belief that heat production is determined by body surface 
is not strictly accurate; a careful analysis of metabolism measure¬ 
ments shows that the heat output is not proportional to the body 
surface. Other factors are the proportion of inert fat and active 
protoplasmic tissue in the body composition, and also the height 
and age. Finally, there are large variations from day to day in 
the same individual. Summing up his conclusion, Benedict says: 

It is clear that the basal metabolism of an individual is a function first of the 
total mass of active protoplasmic tissue and second of the stimulus to cellular 
activity at the time the measurement of the metabolism is made. Apparently at 
present no law can be laid down that will cover both of these important variables 
in the basal metabolism of an individual. 

In the light of this work it is at once seen that units for 
comparing the metabolism are still only in the making, and a 
comparison is only possible of the averages of experiments on large 
numbers of individuals, extended over a long period of time, so as 
to eliminate changes which might be due to the temporary 
ascendency of any one factor. It is obvious, therefore, that 
researches which have been made hitherto on total metabolism have 
not been on a sufficiently large scale to justify any definite 
conclusion, and probably the discrepancies in the results of different 
experimenters may be accounted for in this way. 

The researches into the total metabolism and heat production 
in man have so far not revealed any changes which might be 
attributed to the influence of climatic conditions. It has, however, 
repeatedly been pointed out that the dietary of the majority of the 
aboriginal races in the tropics contains a relatively small proportion 
of protein. This fact has naturally led to the presumption that 
dwellers in the tropics require a less amount of protein in their food. 



3«7 


Protein differs from the other food constituents in the respect that 
it is utilised by the organism in both directions, namely, for 
producing energy and building up tissue. It is, therefore, not 
unexpected that the protein requirements of tropical inhabitants 
has been studied in detail. 

Eijkman (1893) in Java found that Malays consumed 73 3 
grammes of protein per day, of which about 56 grammes were 
actually metabolised, an amount far below the standard usually 
accepted in Europe (100 grammes). An extensive series of 
observations on the protein content of the dietary of native races 
in India has been carried out by McCay (1912), who investigated 
its bearing upon the physique and general efficiency of his subjects. 
In his opinion the small quantity of protein generally consumed by 
the natives was not the result of want of desire, but of inability to 
procure a larger quantity, and he convinced himself that most 
natives would eat more if they could obtain it. He concluded that 
those races with a higher level of protein interchange were generally 
more robust, energetic, and more manly. In the same way the 
daily food of Filipino prisoners, according to Aron (1909), 
contained only 75 grammes of protein. 

A much higher quantity of protein was utilised, according to 
Eijkman, by Europeans living in Java. He found that the food of 
his experimental subjects contained 99*6 grammes of protein, of 
which 88*2 grammes were actually absorbed, which amount 
corresponds to the normal protein requirements in Europe. Other 
observers who examined their food during a visit to the tropical 
zone, confirmed this. Thus Ranke, during his voyage to Brazil, 
found very little change in the protein of his food, and Jaffe and 
Schilling found that nitrogen equilibrium was maintained on the 
same amount of protein in Europe and in West Africa. 

Unfortunately the whole question of the protein requirement of 
man is still unsolved. Atwater’s standard of 125 grammes of 
protein a day, and Chittenden’s standard of about 60 grammes, 
are both accepted by various workers, and recent experiments of 
Hindhede have shown that nitrogen equilibrium can easily be 
maintained upon a much lower amount of protein, provided that 
the diet contains sufficient calorific value. He himself maintained 
nitrogen equilibrium on 32 grammes of protein for 150 days. The 



3*8 


question is further complicated by the necessity of accessory food 
products and their relation to the different constituents of the food, 
about which we are as yet in almost complete ignorance. 

The protein metabolism has been further studied by the 
examination of the various constituents of the urine, since the final 
products of its degradation are excreted in the urine. Eijkman 
found that the urine of Europeans living in the East Indies did not 
differ appreciably from standards in Europe. He found the average 
volume for twenty-four hours as 1,442 c.cm., the specific gravity 
1*017, whilst the average nitrogen content was 13 04 grammes. In 
the physiological text-books the average for Europe is generally 
given as 1,200 to 1,500 c.cm., with a specific gravity of 1015 to 
1*025, and a total nitrogen content of 14 to 18 grammes. He 
allowed for 1 *6 grammes of nitrogen lost in the sweat, and concluded 
therefore that an acclimatized European passed as much nitrogen 
in his urine as he did in Europe. 

Other observers, such as Wick, Ranke and Neuhauss, however, 
found that the urine is much more concentrated. Observations in 
this respect, extending over some time, have been made in 
Townsville by Young (1919). A number of urine samples collected 
during the hot months have been examined in this Institute, and the 
average figures were obtained for the daily urine of twenty-five 
persons, some of whom were engaged in manual labour, and others 
followed a sedentary occupation. The averages calculated are 


given in the Table IV, together with European standards: 

'Fable IV. 


In Townsville 

1 

European standard 

Quantity . 

782 c.cm. 

1,500 c.cm. 

Specific Gravity . 

1.025. 

1.015 to 1-020 

Total Nitrogen. 

10.4 grm. 

16 grm. 

Sodium Chloride . 

1 7.00 grm. 

15 g™ 1 * 

Phosphates . 

173 grm. 

2 to 3.5 grm. 

Freezing Point. 

—0.935 0 to —2.259 0 C. 

—0.87° to —2.71 0 C. 












3«9 


These observations reveal decided differences. The volume was 
much lower and the specific gravity higher. The total nitrogen, 
too, was smaller. The most striking difference lies in the quantity 
of sodium chloride, which may be accounted for by the increased 
quantity of sweat lost, which carries with it sodium chloride. 
Estimations of the amount of water lost from the skin during 
exercise in Townsville have yielded interesting figures. After a 
brisk w’alk of about seventy minutes with an external temperature 
of 24'4°C. (76° F.) wet bulb, as much as 1,100 grammes in weight 
have been lost. As the subjects of these experiments were weighed 
in their clothing, this figure only represents the water which had 
actually evaporated. Samples of sweat carefully collected contained 
from o* 1 to 0*3 gramme of sodium chloride per 100 c.cm., and the 
cutaneous excretion would account for several grammes of sodium 
chloride in the above experiment. The nitrogen lost in the sweat 
can only be small, and is almost negligible, since samples of sweat 
only contained 0 03 to 0 04 grammes per 100 c.cm. 

The urine measured during the cold weather averaged a higher 
volume, 1,357 c.cm., a lower specific gravity, 1019, and an 
increased salt content, 9 44 grammes, per day. 

The total nitrogen in the urine in Northern Australia was 
decidedly below the European standard, even considering that a 
small amount (1 to 2 grammes per twenty-four hours) may leave 
the organism by way of the skin. There is thus an indication of a 
decrease in the amount of protein metabolised. Whether this is 
due to a smaller intake of food brought about by a want of appetite 
during the hot weather, so commonly observed, or whether other 
factors come into play, must be left an open question. 

(/) Effect of a Tropical Climate on the Nervous System . In 
many of the writings on health and disease in the tropics attention 
has been drawn to the frequent occurrence of a mental condition 
resembling neurasthenia, of the same type as that found in 
temperate climates. It occurs mostly amongst Europeans, women 
and men, after a prolonged residence in the tropics, and is referred 
to in the literature as tropical neurasthenia. In its mildest form it 
manifests itself in a lability of the mental equilibrium, fits of 
depression alternate with states of exuberance; unwarranted 
irritability over trifling matters is hardly ever absent, leading to 



390 


uncontrollable outbreaks of temper. It is associated with a loss of 
mental activity and power of concentration, lack of confidence and 
failing memory, all of which cause a decreased working capacity. 
The realisation of this state, and the futile attempts to overcome it 
by increased spurts of energy, tend to exaggerate the trouble. This 
condition may be stationary for a long time, influencing the career 
of the sufferer, or may get steadily worse and lead to a condition 
termed by Plehn 'tropical fury’ (Ttopen Koiler), which often 
explains the committal of deeds of violence in an outburst of 
uncontrollable passion. 

Amongst others, Fales (1907) has drawn attention to the 
occurrence of neurasthenia amongst men and women after a 
residence of a year or more in the Philippines, which in women is 
often associated with disturbance of menstruation and of the 
vasomotor system. In his opinion it is in most instances not directly 
attributable to parasitic invasions, but to climate only, and forms 
one of the chief drawbacks to acclimatization. Woodruff made the 
same observation in the same locality, and attributed the condition 
entirely to the effects of tropical light. 

In 1913 the Society of Tropical Medicine devoted a meeting to 
the discussion of this subject, at which the President, Sir Havelock 
Charles, delivered an introductory address on * Neurasthenia and its 
Bearing on the Decay of Northern Peoples in India/ He discussed 
the occurrence of * Punjab head * in Bengal, which is characterised 
by shortness of temper, forgetfulness, sleeplessness and disinclina¬ 
tion to work, etc., which corresponds to tropical neurasthenia. 
Similar 'heads 1 occur throughout India and other parts of the 
tropics, and affect Europeans after having lived in the tropics for 
a considerable period without furlough. In a number of cases, 
according to Havelock Charles, tropical disease may be held 
primarily responsible for the condition, but in many instances he 
considers climate alone as the root of the evil. Since ‘the constitu¬ 
tion of the northern races is developed in temperate latitudes/ he 
believes 'that its powers are injuriously affected by the climatic 
conditions of the hot zone, and this must be attributed to damage 
done mainly to the nervous system by a hot and humid climate/ 
Unfortunately his address, and a great deal of the discussion which 
followed, were marred by statements which were merely expressions 



39 1 


of personal opinions, collected during a shorter or longer residence 
in the tropics, and often coloured by prejudice; actual data in 
support of these opinions were sadly lacking. 

Basil Price (1913), who for many years examined the causes of 
invaliding home of members of the Church Missionary Society, 
found that neurasthenic conditions were the cause of 25 per cent, 
of invalidity in most countries. 

A perusal of the general literature on the tropics confirms the 
fact that a complaint similar to neurasthenia is very prevalent 
amongst Europeans who have emigrated to the tropics, but it is 
difficult to gauge how far climatic influences alone may be held 
responsible, or what part is played by the altered conditions and 
habits of tropical life. The novelty of the strange environment 
often leads to an increased output of energy, and it is only after 
a time that the newcomer realises his energy capacity under the new 
conditions and begins to husband his strength, but often too late 
to avoid paying the penalty. 

It is, however, beyond doubt that the monotony and discomfort 
of life and climate, the lack of pleasure and excitement, the big 
distances from the centres of civilisation and, in women especially, 
the confinement to the house during the hot hours of the day, all 
help towards producing a mental condition resembling neurasthenia. 

(g) Clothing in the Tropics. The degree of bodily comfort in 
the tropics, in the same way as in a cold climate, is governed by a 
number of factors, many of which may be modified artificially. In 
a cold climate bodily comfort is attained by conservation of heat, 
in a hot climate by the allowance of a maximum amount of cooling, 
and the clothing is a very important factor in this respect. 

The literature contains numerous accounts of experiments and 
suggestions on the most suitable kind of clothing for a tropical 
climate. The aboriginal inhabitants possess a dark pigmented 
skin, and, theoretically, therefore, a dark-noloured garment, 
allowing of ample ventilation, would appear to be the ideal, since 
it would imitate nature. Practice, however, does not bear out this 
assumption. It has been shown that a dark skin absorbs heat rays 
to a greater extent than does the white skin; its temperature, 
therefore, rises more quickly and the sweating mechanism is 
brought into action earlier and the cooling due to evaporation is 



39 2 


more effective. A dark dress material absorbs heat in the same way 
as a dark skin, promotes perspiration, but at the same time acts 
as an obstacle to the free evaporation of sweat from the skin. There 
is between the skin and the clothing a space filled with stagnant 
air, which in fact contains more moisture than the surrounding air, 
on account of insufficient ventilation. In consequence, the dress 
material becomes more and more impregnated with moisture, and 
the meshes of the fabric clogged with water, and the degree of 
saturation of the air space between skin and clothing exaggerated. 
*Young has recorded observations in North Queensland upon the 
temperature between skin and shirt—‘skin-shirt temperature'— 
which confirm and amplify Hill's (1914) previous experiments in 
the same direction, which were carried out in England. Wet and 
dry buib thermometers were used, the bulbs, of which were encased 
in wire cages to prevent contact. The temperatures thus observed 
were invariably several degrees higher than the outer temperatures. 
Hill's observations, made during an English winter, showed figures 
as high as those recorded in North Queensland, but his subjects 
wore heavy clothing suitable to winter conditions in Europe, 
whereas Young's subjects only wore a thiji shirt. This is a clear 
proof that the quantity of clothing and the nature of the material 
is of paramount importance in determining the condition of the air 
surrounding the skin. The former does not require any further 
explanation; the latter, however, the nature of the material, may 
be further discussed here. It should possess two properties, firstly, 
that of absorbing moisture and allowing of a maximum surface for 
speedy evaporation—the difference between woollen, cotton and 
silk material is well known—and, secondly, it should prevent, as 
far as possible, penetration of heat waves. 

Attempts to obtain this objective have been made by several 
workers, who experimented on the suitability of material of various 
colours for tropical use. This question has been discussed by 
Sambon (1907), who based suggestions upon experiments carried 
out by Baly upon the penetration of rays from an electric arc 
through a piece of dark pigmented skin derived from an Indian. 
Baly photographed the spectrum after passage of the rays through 


* See these Annals, p. 313. 



393 


the skin, and noticed that rays of shorter wave length than 3 ,6 ooju 
(ultra-violet) were entirely absorbed, proving thus that the skin 
pigment excludes the ultra-violet rays. From these observations 
Sambon suggested that, for tropical use, material coloured black, 
red or orange should be used, colours which eliminate the rays of 
short wave length. As all these colours absorb heat rays, he 
suggested at first a combination of white outer garments and 
coloured underwear, the former to reflect heat rays, the latter to 
exclude actinic rays. Later he suggested the manufacture of a 
special material, combining both advantages, and this has been put 
on the market as ‘Solaro.’ This fabric is composed of white and 
suitably coloured threads, woven in such a way as to present a warp 
(upper surface) of white, and a weft (under surface) of orange, red 
or black, and it has been proved by spectroscopical examination to 
be as impervious to actinic rays as the black skin. 

Schmidt (1909) suggested the wearing of loose garments made 
of either thick light-coloured or thin dark-coloured material, and 
recommended the use of material similar to that suggested by 
Sambon. 

The practical test of coloured underwear, however, has not 
proved it to be advantageous. Phalen (1910) conducted experi¬ 
ments on a large scale on soldiers in the Philippines. He supplied 
five hundred soldiers with orange-red underwear, and compared 
their well-being in the course of a year with another group wearing 
white undewear. He compared body weight, the number of red 
cells and haemoglobin, blood pressure, pulse and respiration rate, 
body temperature, and incidence of sickness in both groups. He 
found that the men wearing orange-red underwear showed changes 
due to heat, such as loss in body weight and haemoglobin and 
decreased blood pressure, more marked than the control group. 
Phalen concluded from his experiments that the coloured garments 
were more receptive to heat rays than were the white, since wearers 
complained persistently of greater heat, greater weight, and 
increased perspiration, and expressed the opinion that khaki 
clothing alone affords the same protection from chemical rays as 
the special underwear. 

The question of clothing in the tropics has been dealt with from 
a more commonsense point of view by Gibbs (1917). ‘Clothing 



394 


which in the sun will cast a shade upon the body without hindering 
the air circulation and heat radiation will be the most desirable, 
and if a colour is used which will give a minimum of heat absorption 
the efficiency is increased. This ideal condition is fulfilled by the 
umbrella, and it is evident that a large white umbrella lined with 
a material of a colour agreeable to the eyes, for example, a shade 
of green, will be most efficient/ He states that, since custom 
prescribes that man shall cover his body, this mode of * clothing * 
is in fact impracticable, and substitutes for the umbrella a large- 
brimmed helmet, casting a shadow over the back, shoulders and 
chest, and a loosely fitting white suit of material as thin as possible. 

Judging by our own experience, the wearing of light-coloured 
suits of porous material assures the maximum amount of comfort 
possible, but it is obviously impossible for a labourer to wear white 
clothing at his work. In North Queensland he wears, as a rule, 
khaki-coloured trousers and a thick, dark-coloured flannel shirt, 
which prevents the sun’s rays from penetrating to the skin, and 
absorbs perspiration, and thus allows a maximum degree of 
evaporation of the sweat. 

On the whole, an extended inquiry amongst the residents in 
tropical Queensland has revealed such a diversity of opinions as to 
what type of clothing is the most comfortable that it appears 
presumptuous to lay down definite rules. Many prefer cotton for 
underwear, others pin their faith on wool, but all agree on the one 
point that outer as well as under garments should be as porous and 
as light in weight as possible. 

(3) Summary of Physiological Changes of the White Man in the Tropics 

A review of the known facts concerning the physiological changes 
of the white man transplanted to the tropics is, from a scientific 
point of view, disappointing. Throughout the literature published 
since the beginning of the last century may scrappy attempts have 
been made to obtain evidence in this respect; figures have been 
collected and experiments on metabolism and other physiological 
functions have been performed. In spite of the energy expended, 
the results are not conclusive; firstly, on account of the smallness 
of the number of observations recorded by the different individual 
observers, too small to eliminate individual variation and error, 



395 


secondly, on account of the uncertainty of the generally accepted 
standards which may be considered normal for a temperate climate. 
The question of body temperature' is a case in point. Several 
observers formed their conclusions by accepting 37 0 C. (98'6° F.) 
as a normal body temperature, and considering even a small rise 
above this as an indication of an increase in body temperature in 
the tropics. If a series of observations had been taken on a few 
subjects at intervals during the day and night, and the average 
calculated, discordant results would have been the outcome, even 
in Europe. 

The same criticism applies to experiments regarding other 
physiological functions, as, for example, the total metabolism as 
measured by the gaseous exchange. As already pointed out, a 
great deal of work has been attempted in Europe, and recently on 
a much larger scale than hitherto in the large nutrition laboratories 
in the United States. In spite of these concentrated and intensive 
efforts, as Benedict has pointed out, no definite unit for the 
comparison of the metabolism of normal individuals has been 
discovered, and there does not exist at present a definite knowledge 
of the factors influencing individual variations. 

Such changes as have been noted have been of a qualitative, 
but not of a quantitative, nature. Thus, with regard to maintenance 
of body temperature under tropical conditions, no evidence could 
be found in support of the view that heat, generated by internal 
combustion, is reduced in order to regulate the body temperature; 
even under the altered conditions the body still relies on physical 
means for this purpose. As convection and radiation of the body 
are greatly decreased with the high temperature, cooling must be 
brought about to a much larger extent by sweating and evaporation 
than in a temperate climate. 

Again, the increased loss of fluid from the body by the skin is 
accompanied by a decrease in the quantity of the urine voided. As 
the Sweat contains sodium, chloride, an increased amount of this 
salt must be lost through the skin, which in its turn again leads to 
a decreased amount of salt excreted in the urine. 

Blood examinations performed on a large scale have not proved 
the occurrence of a tropical anaemia as such, since the figures for 
haemoglobin and for the formed elements do not show a definite 



decrease. There is a certain alteration in the percentage of the 
various types of white blood corpuscles, and in the number of cell 
fragments in the polymorphonuclear neutrophile leucocytes, showing 
that the percentage of ‘ young leucocytes ’ in the circulation, that is, 
leu9ocytes containing a smaller number of nuclei or nuclear 
fragments, is larger than in a temperate climate. This is expressed 
in hacmatological language as a ‘ shift of the Ameth index to the 
left.’ The significance of this change is, however, not dear, but at 
any rate does not seem to indicate a decreased power for resisting 
disease, as supposed by the Philippine observers. 

The concensus of opinion gleaned from experiences in various 
parts of the tropics seems to indicate that living in the tropics affects 
the nervous system, and that neurasthenia seems to be more 
prevalent than in a temperate dimate. It is, however, impossible 
to obtain definite figures and data, and most statements are only 
based on personal experience. In the light of this it would be 
interesting to ascertain whether a detailed examination of the 
functions of the nervous system would reveal any definite alterations, 
which, on account of their frequent incidence, would have to be 
considered an outcome of life in a tropical climate. 

It appears, therefore, possible that a closer study of the 
functions of central and peripheral nervous system might reveal 
definite alterations. It would be advisable to obtain figures for the 
promptitude and quality of the response given by the nervous 
system to various stimuli. Reaction times and responses to different 
stimuli are quantities readily capable of exact measurements by 
means of generally used apparatus. 

An extensive investigation into the mental activities in general 
would perhaps yield figures of definite value. The testing of a 
great number of individuals in the tropics on similar lines to those 
applied in choosing aviators would be very interesting. The 
applying of the Binet-Simon test for mentality to a great number 
of school children, again, would furnish figures of definite value for 
deciding the presence or absence of racial degeneration. In the 
mentality tests the methods are rough, and only a very large 
number of tests, sufficiently large to eliminate personal error on the 
part of the observer and to minimise the influence of a few feeble¬ 
minded children, would yield figures of value. In short, all the 



$97 

efforts to detect physiological changes due to living in a tropical 
climate have not yielded definite results. It would, however, be 
premature to conclude that such changes do not exist, and it is 
possible that the advent of refined methods and more sensitive 
apparatus may in the future demonstrate definite alterations. 


WHITE SETTLEMENT IN TROPICAL AUSTRALIA 

(i) General Conditions and Statistics 

When comparing tropical Australia with other countries situated 
in the torrid zone, it becomes apparent that Northern Australia 
occupies a somewhat unique position from more than one point of 
view. On the whole, the general conditions of a tropical country 
are modified to a much greater extent by climate -per se, meaning 
seasonal incidence and degree of rainfall, than a country within the 
temperate zone. With the exception of a wet belt on the Queensland 
coagt, mentioned previously, the rainfall is limited to a few months 
in the year, and during the remainder of the year at the most an 
occasional light shower disturbs the monotony of sunshine, and 
during about seven months of the year the weather conditions 
correspond to the ‘dry tropics.’ Vegetation, which is luxuriant 
during the wet season, dies down, and the green of the countryside 
gives place to a uniform brown. The shrivelled up undergrowth 
is often consumed by bush fires, which are said to be sometimes 
started by an accidental focussing of the sun’s rays, and which 
sweep for miles over the country, leaving only the trees standing, 
bare of their bark. 

Many phases of insect life, which are dependent on vegetation 
and moisture, die down during the dry months, only to become 
all-pervading again as soon as the rains start. These changes are 
more marked inland than on the coast, since many of the inland 
districts have a very small rainfall and the hot winds arising from 
the barren desert of Central Australia cause the inland plains of 
Northern Australia to approximate to a typical ' desert type ’ of 
climate, with cold nights and scorching hot day 9 . 

Another respect in which Northern Australia differs from other 
tropical regions is the spardity of the aboriginal population. The 



398 

natural conditions of Northern Australia seem to militate against a 
largo aboriginal population, and the natives have never evolved 
beyond the nomad state and have for an unknown reason never 
made any attempt towards settlement. The uncertain food 
conditions, aggravated by droughts, hive helped to keep down 
their numbers. After the arrival of Europeans, the inability of the 
aboriginals to change their nomadic habits has led to a decrease in 
their number and has prevented them from living alongside the 
white man. In consequence, in any part where a large white 
population exists, the black man has become extinct. 

The white population is still very scanty, and is mostly 
concentrated in a few towns which are long distances apart. 

The climatic conditions, especially the small and erratic seasonal 
rainfall, in conjunction with the scarcity of population, have created 
the unique position which Northern Australia holds to-day. The 
long dry season influences the insect life, and those tropical diseases 
which are transmitted by certain insects, such as the mosquito, are 
apt to show a corresponding seasonal incidence, and are to a certain 
extent kept in check. Other diseases, such as infectious and 
contagious ailments, are in a similar way kept down by the 
sterilizing action of the sunlight, which may kill the virus before it 
can spread over large and scantily populated areas. 

The contention that the prevalence of certain diseases is favoured 
by larger and more evenly distributed rainfall is borne out by the 
fact that in districts situated within the wet belt a greater incidence 
of certain diseases is observed. Various fevers, such as endemic 
glandular fever, other scrub fevers of unknown aetiology, which are 
most probably insect-borne, occur throughout the wet belt, but are 
absent from other drier districts. In the same way hook-worm 
disease is far more prevalent in those places where the rainfall is 
higher, and is, as far as our present experience goes, practically 
absent from other districts showing similar local conditions but a 
smaller and strictly seasonal rainfall. 

The effects of climate and surroundings upon any race Ends a 
clear expression in the vital statistics, such as birth and death rates, 
infantile mortality rates, and expectation of life. 

The statistical data for North Queensland and Northern 
Australia have not been published separately for comparison with 




399 


similar statistics for the rest of Australia; moreover, it is impossible 
to obtain definite data as to the number of inhabitants in Australia, 
since the last census took place in 1911, and the population of 
Northern Australia Is of a migratory character; any figures obtained 
are therefore only of approximate value. The Commonwealth 
Statistician, Mr. G. H. Knibbs, C.M.G., was able to supply 
information which throws light upon this question, and has kindly 
given permission to include his statements in the present publication. 

The area and population of the tropical portion of the Common¬ 
wealth are shown in the following table: 


Tabu V. 


Area and population of the tropical portion of Australia. 


Tropical Portion of— 

Area in 
square miles 

Popul 

a 

ation (exclusive of full-b 
boriginals) at Census of 

ilooded 

Northern Territory . 

Queensland . 

Western Australia . 

426,320 

359.«» 

364,000 

1881 

3f45 f 

56,041 

661 

1891 

4,898 

108,986 

3,7” 

1901 

♦,096 

>45.9** 

4,664 

1911 

3.3>° 

157,112 

4,99* 

Total . 

1,149,320 

60,153 

**7,595 

*54,74* 

165420 


These figures relate to all that portion of the Commonwealth 
which lies to the north of the Tropic of Capricorn. For the tropical 
portions of Queensland and Western Australia population figures 
are available at the date of a census only. For the Northern 
Territory they are compiled quarterly, and the returns for 
December 31, 1917, give a total of 4,908—an increase of 1,598 since 
the census of 1911. 

At the census of 1911 the total population of Queensland 
(exclusive of full-blooded aboriginals) was 605,813, so that at that 
date the tropical population of Queensland represented rather more 
than 25 per cent, of the total. It also represented about 95 per cent, 
of the total tropical population of Australia. It might consequently 
be presumed that any unhealthiness of tropical Australia would have 
left its impress on the vital statistics of Queensland. That this is 
pot the case is clearly shown from the following comparison of the 



400 

Queensland death rates per thousand of population with those for 
the Commonwealth as a whole: 

Tabu VI. 

Comparison of Queensland and Commonwealth Death Rates, 1906 to 1917 



1906 

! 9°7 

1908 

* 9°9 

1910 

1911 

1912 

* 9*3 

1914 

* 9*5 

1916 

1917 

Queensland 

9.50 

10.31 

10.26 

9.70 

971 

10.65 

10.96 

* 0.39 

9-97 

11.00 

11.09 

964 

Commonwealth 

10.92 

10.99 

11.07 

10.33 

io -43 

10.66 

11.23 

O 

00 

10.50 

10.66 

11.04 | 

9.80 


It will be seen that in ten out of the twelve years under review 
Queensland recorded a lighter death rate than the Commonwealth 
as a whole, and that in one of the other two years (viz., 1916) the 
difference was inappreciable. The remaining year (1915) was that 
in which Queensland suffered from the most severe drought ever 
known there, while the greater part of the remaining States 
experienced normal conditions, having suffered from drought in 
the preceding year. 

Another test of salubrity often applied is that derived from a 
comparison of the deaths of infants under one year of age with the 
total births. The rate so deduced (i.e., the number of deaths of 
infants under one year of age per one thousand births) is what is 
generally, known as the ‘ infantile mortality.’ A comparison of the 
Queensland results with those of the Commonwealth as a whole for 
the past twelve years is furnished in the following table: 

Tabu VIL 

Comparison of Queensland and Commonwealth Infantile Mortality, 1906 to 1917 



1906 

I 9°7 

1908 

« 9°9 

1910 

19U 

1912 

1913 

1914 

* 9*5 

1916 

1917 

Queensland 

74.68 

77-65 

70.67 

71.50 

62.90 

65.36 

71-73 

6335 

63.93 

6*.33 

70.27 

S387 

Commonwealth 

83.26 

81.06 

77 - 7 * 

71.56 

5 

bo 

68.49 

7**74 

72.21 

7«-47 

67.52 

70.33 

55 - 9 * 


In every one of the twelve years under review the infantile 
mortality of Queensland was more favourable than that of the 
Commonwealth as a whole. Further, in both the foregoing tables 
the comparison has been between Queensland on the one hand and 
the Commonwealth, inclusive of Queensland, on. the other. A 
comparison between Queensland and the Commonwealth exclusive 
of Queensland would have given results even more favourable to 
Queensland than those deduced above. 




4oi 


In the absence of complete data in respect to the population and 
mortality of the tropical parts of Australia, it is impracticable to 
carry the test further, but the figures given above indicate that the 
vital statistics of the Commonwealth furnish no evidence of lack of 
salubrity in those parts. On the contrary, the State having 25 per 
cent, of its population within the tropics, and containing some 
95 per cent, of the tropical population of the Commonwealth, has 
a record for general and infant mortality much more favourable 
them that for the Commonwealth as a whole. It may be noted that 
practically the whole of the State of Queensland lies in the north 
of the twenty-ninth parallel of south latitude. 

Striking as the foregoing Queensland figures are in comparison 
with those for the whole of Australia, an even more telling result 
is obtained by comparing them with those of some of the leading 
European and other countries. 


Table VIII. 


Country 

Year 

Death Rate 

Rate of 
Infantile 
Mortality 

Queensland . 

1913 

10.4 

63 

Commonwealth . 

1913 

10.8] 

7 * 

Netherlands . 

1913 

12.3 

9 * 

Denmark. 

■ 9*3 

12.5 

94 

Ontario (Canada). 

1913 

12.7 

* *7 

Norway . 

* 9*3 

*3.2 

65 

Sweden . 

* 9*3 

13.6 

7 ° 

England and Wales . 

* 9*3 

13.8 

108 

U.S.A. (registration area) . 

* 9*3 

* 4 * 

• 

Switzerland . 

* 9*3 

* 4-3 

96 

Belgium . 

* 9*3 

14.8 

120 

Germany. 

* 9*3 

*5.0 

* 5 * 

Scotland . 

* 9*3 

* 5-5 

no 

Ireland . 

* 9*3 

* 7 * 

97 

France . 

* 9*3 

* 7-7 

98 


Not available. 






















402 

Table VHI—conti*iud 


Country 

Year 

Rate 

Rate of 
Infantile 
Mortality 

Italy . 

1913 

187 

*37 

Japan . 

* 9*3 

* 9-5 

*50 

Austria . 

1913 

20.5 

1S0 

Jamaica . 

1913 

21.7 

* 7 * 

Spain . 

* 9*3 

22.1 

• 

Hungary . 

* 9*3 

23.3 

186 

Rumania.. 

* 9*3 

2S.9 

202 

Ceylon . 

* 9*3 

28.4 

189 

Chile . 

* 9*3 

30.1 

2 55 


* Not available. 


In addition to the foregoing, it may be pointed out that the 
moitality experience of Queensland has been continuously improving, 
both absolutely and also in relation to the experience of the 
Commonwealth as a whole. This is clearly brought out. in the next 
table, showing the expectation of life at age o in each of the last 
three decades: 


Table IX. 

Expectation or Lir* at Age o. 



Males 

Females 

_L 

Decade 

Queensland 

C’wealth 

Queensland 

C’wealth 

1881-1890 .. 

Years 

4 ** 33 ° 

Yean 

4 r «99 

Years 

49*754 

Years 

50-844 

1891-1900. 

49-5*2 

51-076 

55-80° 

54756 

1901-1910. 

54*203 

55.200 

59' 2 94 

58-837 


In Mr. Knibbs’s opinion, local statistics relating to individual 
towns would not give results sufficiently reliable to warrant definite 
conclusions, owing to the migratory character of much of the 
population, and, in connection with birth, owing to the tendency to 
move into the towns for purposes of confinement. 




























4°3 


(2) Housing 

The question of the construction of suitable dwellings in the 
tropics has been the object of a great deal of study and controversy. 
A suitable tropical residence should be so constructed that the 
interior is protected as much as possible from the direct rays of the 
sun, in order to prevent excessive heating of the walls of the room, 
and at the same time sufficient ventilation should be provided in 
order to secure the quickest possible cooling of the structure. 
Unfortunately, in the construction of the average dwelling house in 
North Queensland these principles have not been sufficiently 
considered. The greater proportion of the dwellings of the settlers 
on the land are entirely unsuitable, and a great number in the larger 
towns are far from ideal. ' A statement of a leading firm of 
architects with twenty years’ experience in the north is of interest, 
namely, that they have never designed one cottage as they know a 
cottage should be designed, on account of the prejudice against the 
introduction of novel ideas. 

The majority of the scattered habitations of the smaller settlers 
in north-western Queensland are built of galvanized iron on a 
wooden framework, without verandahs or any insulation, and the 
temperature of the interior during the hot hours of the day surpasses 
by far the outer air temperature. 

The small dwellings in the towns are raised on piles, and 
contain as a rule four rooms; they possess a narrow verandah in 
front, and sometimes also on the side. The rooms are as a rule low, 
and the roof is composed of galvanized iron sheets and often does 
not possess an inner lining. The kitchen is nearly always detached 
and generally consists of a small cubicle built of galvanized iron 
only. Most of the larger wooden houses have a verandah sill round, 
and higher rooms with ample ventilation. 

A good type of tropical house may be seen in Darwin, where 
many of the houses are constructed after a design similar to houses 
in the Far East. The verandahs are wide and are closed in by 
bamboo shutters, which are kept shut during the heat of the day 
and thus prevent the heating up of the rooms and at the same time 
allow of sufficient ventilation; the shutters are opened as soon as 
the sun gets low, 



It is quite evident that a galvanized iron ‘humpy/ without 
verandah, is the most unsuitable structure to reside in in a hot 
climate. During the day the heat is nearly unbearable, and the 
only advantage is the speedy cooling of the walls after sundown. 
The larger houses, built of timber, with open verandahs, are, on 
the whole, suitable for the climatic conditions if attention has been 
paid to several points. 

The house should be raised off the ground and built on piles in 
order that the air should have free access to every part of the house. 
It should be so placed, if practicable, that it lies in the direction of 
the prevailing breeze, and windows and doors should as far as 
possible be opposite, in order to allow of the maximum of ventila¬ 
tion. The verandah should be sufficiently broad to protect the walls 
from the direct rays of the sun at any time, but the minimum width 
depends upon the latitude in the geographical sense. According 
to Schilling (1909), the houses in regions near to the equator require 
a verandah on the four sides, as the path of the sun is more vertical. 
In regions below the Tropic of Capricorn a verandah on the south 
side is not absolutely necessary, as the path of the sun is inclined 
from the north and the south side of the house is not struck by the 
direct rays at any time. The rooms should be spacious and with 
as many windows and doors as practical for the sake of ventilation 
and lighting. It is only too well known that mosquitoes and other 
insects choose dark comers and avoid light. A roof of galvanized 
iron is suitable provided that the necessary ventilation is allowed 
for. The roof should be slightly raised from the wall.* The top 
gable should be provided with a ventilator and, if possible, there 
should be louvres on either side below the gable to provide 
additional ventilation. If the roof be closed in, there is a layer of 
hot, stagnant air between the roof and the ceilings of the rooms, 
which is heated by the sun to a higher temperature than the outside 
air, and this keeps the temperature of the rooms high after stinset. 

We are indebted to Messrs. Lynch and Hunt,- architects, of 
Townsville, whose activities extend throughout North Queensland, 
for the framing of the following recommendations for building small 
dwellings suitable for North Queensland, and costing approximately 
from £200 to ;£6oo: 

Apart from any further items that may be found advisable, as many of the 



4os 


following recommendations as the site will permit should be embodied in every 
cottage:— 

1. Buildings should face due east and have verandahs to front and back of 
not less than nine feet in width. 

2. In addition to other necessary openings, there should also be doors and 
windows so arranged in every room to allow the wind to enter on the weather 
6ide and escape on the lee side. Doors should also be arranged so as to avoid the 
necessity of retracing one’s steps to get to any section of the house. 

3. Buildings should not be more than one room in depth. 

4. The size of a room built under the most favourable circumstances should 
be regulated by the number of prospective occupants. Eight hundred cubic feet 
per head would be a fair minimum. 

5. Side verandahs and excess widths to other verandahs should not be con¬ 
structed at the expense of the size of the rooms. Large rooms and limited verandah 
space are infinitely better than small rooms and wide verandahs. 

6. Blocks should be approximately 2 feet 6 inches above the ground. They 
should be either brick or concrete. 

7. All buildings should be constructed to resist cyclones. 

8. Fully exposed walls should be double-sheeted and ventilated. 

9. Roofs should be fitted with ventilators. 

10. Ceilings should stand two feet below the top of top plates. In this con¬ 
nexion studding should not be less than twelve feet in length for the smallest 
cottage and proportionately longer for larger cottages. 

11. Fan-lights should be hung immediately under ceilings. 

12. Ledges and corners should be avoided as far as possible. 

13. Roofs, should be hipped and continuous, having eaves approximately two 
feet in width, to walls and verandahs. 

Mr. C. D. Lynch has kindly put at our disposal a novel 
suggestion for building suitable houses, which would entail an 
expenditure of at least a thousand pounds, and which scheme is 
the outcome of twenty years’ practical experience of house designing 
in North Queensland. The building, the idea of which is shown 
in the accompanying sketch (fig. VI), is constructed of reinforced 
concrete or brick, and possesses double outer walls, with an 
intervening and suitably ventilated air space of about three inches, 
thus ensuring coolness and dispensing with the necessity of 
verandahs. The floor space is sub-divided by wide corridors 
(3*5 to 4*5 metres, or about 12 to 15 feet) arranged in the form of a 
cross, on to which the rooms, placed at the comers of the house, 
open. The corridor may serve as a general living room, since it 
enjoys the maximum amount of ventilation, whatever the direction 
of the wind may be. A flat roof, surrounded by a balustrade, 
would, according to Mr. Lynch, add further to the roominess and 
comfort of the building. 

Suggestions are found in the literature for cooling houses by 



406 

artificial means. Attempts have been made to circulate artificially 
cooled air through buildings, but in practice this method has not 
proved satisfactory on account of the high initial expense of the 
necessary machinery and the technical skill required for its main¬ 
tenance. Less costly, but at the same time less effective means, arc 
fans run by electrical or water power; but as the majority of the 
northern towns of Queensland are without electrical power, and as 
the water rate is excessive and the supply often uncertain, fans in 
private houses have not come into general use. 



/Jf -/ —/* 


Fig. 6. 

(Reduced one half from original drawing.) 

The systematic laying out of towns in tropical Australia has 
unfortunately been sorely neglected, and most settlements have been 
allowed to grow up in a haphazard way. In the' main streets of 
the larger towns ground has become too valuable to permit of an 
adequate space between buildings to allow free circulation of air. 
The main streets have often been laid out regardless of the 
prevailing winds. Flinders Street, the main street in Townsville, 
runs in the direction from north to south, the prevailing breezes 
blowing from south-east, and, as a result, the street has the well- 




4 °7 

deserved reputation of being the hottest comer in North Queensland. 
If, in the original laying out of the town, this point had been 
considered and a number of cross streets in the direction of the 
prevailing winds allowed for, a great deal of discomfort might have 
been spared. 


GENERAL RtSUMi 

A consideration of the position in Northern Australia at present 
shows evidence that the progress in North Queensland, and to a 
still greater extent in the Northern Territory, has not been 
commensurate with the advance of time and with the undeveloped 
resources of the country. Certain townships, without doubt, have 
gone ahead, have increased in population and prosperity, but at 
the same time other towns have ceased to prosper, have decreased 
in population, and their inhabitants have migrated to the prospering 
townships. It is, however, difficult to get definite figures which 
would illustrate this point, as the census is only taken every ten 
years, but the relatively small increase of the population of the 
north when compared with the south of Queensland during the 
decade between two census (1901-1911) is significant. The total 
increase in population for Queensland during this time amounted 
to 108,657 souls; of these, only 7,577 represent the increase in the 
north. Considering that in 1911 25 per cent, of the total popula¬ 
tion of Queensland was living in the northern division, the increase 
in district was relatively much smaller than in the centre or in the 
south. Even if figures for the last seven years could be obtained, 
their value for gauging the progress of the country would be 
doubtful, on account of conditions brought about by the war which 
led to a nearly complete cessation of immigration. 

The population of the Northern Territory has slightly increased 
lately. Even so, at the end of 1917 the total population is 4,908, 
and represents approximately one inhabitant to about 80 square 
miles. 

It is therefore obvious that Northern Australia is not as favoured 
for settlement as the southern parts of the Commonwealth, and in 
this connection it was pointed out by Sir Thomas Anderson Stuart 
on the occasion of a meeting of the Royal Colonial Institute, in 1912, 



that the southern and more salubrious parts of Australia were not 
by any means over-populated yet, and that immigrants naturally 
preferred to settle there than to go further afield; thus, in the course 
of time they would find it an easier matter to obtain a settlement 
in the northern parts than in the south. It was simply a case of 
filling up, and the rapidity would depend on the rate of immigra¬ 
tion. A similar conclusion was arrived at by Atlee Hunt (1915) 
regarding the Northern Territory. ‘ Should any new and rich 
mining field be discovered, the question of populating the Territory 
will settle itself; but in the absence of any such happening, it is 
submitted that the only course open is to wait until, in the natural 
course of things, the trend of population moves gradually 
northward.* 

It is well known that economic conditions play an important 
part in the settlement of any country, and especially of the tropics. 
There is no doubt that life in the tropics is burdened with a great 
deal of discomfort, due to heat and other general conditions brought 
about by climate. For this reason the settlement of the tropics 
carries wdth it more difficulties than settlement in a temperate 
climate. This is borne out in practice by the experience that, 
throughout the world, the white population in tropical parts is 
largely of a migratory character. Tropical Australia does not form 
any exception to this rule. Even during a comparatively short 
residence one notices that the population is continuously changing, 
and that there exists a general desire amongst the inhabitants that 
their stay in the north shall be as short as possible, and the whole 
aspect of the majority^of northern towns bear testimony to this. A 
lack of public interest is noticed everywhere, and has led to putting 
up with makeshifts. Houses are only rarely built for comfort, as 
their owners hope to occupy them only for a few years. Town 
improvements are often only of an ephemeral character, and short¬ 
sighted policies are the rule in most instances. This again is due 
to the unsettled mental condition of a migratory population which 
is unable to concentrate on anything but the most pressing 
immediate needs. 

Many conditions co-operate in bringing about this state of 
affairs. North Queensland is the most recently settled part of 
Australia; it is an enormous stretch of country, with comparatively 



few lines of communication with the centres, and consequently it 
has received a great lack of consideration from the centres of 
government. A feeling of isolation is a natural outcome, and 
exaggerates migratory tendencies and the desire to get back to 
‘ civilisation.’ 

The hot summer naturally militates against permanent settle¬ 
ment, especially since the housing, lack of water, etc., do not 
mitigate the severe discomforts. If a person residing in northern 
Europe were obliged to live during a severe winter in a draughty 
wooden house, without artificial means of heating, a strong desire 
to move to a more congenial climate would be created. In the same 
way one could not expect a white population to thrive in the tropics 
unless all possible means be adopted to alleviate climatic conditions. 

Many newcomers arrive in the north with a prejudice against 
northern Australian heat, which has been instilled into them by 
their southern friends, and adds greatly to this discomfort. Dressed 
in their southern clothing they walk about perspiring, but never 
think that a cold day in the south, spent in a light cotton suit, 
would be equally, if not more, uncomfortable for the opposite 
reason. 

In addition to climatic influences, the mode of employment in 
general tends to attract a migratory population. North Queens¬ 
land is dependent almost entirely on raw products for supporting 
its inhabitants, and permanent institutions, such as factories, giving 
employment throughout the year, are lacking. 

The settler in the north has thus to face conditions which, when 
compared with those of a temperate climate, render life in one way 
more uncomfortable, although in another way less strenuous on 
account of the lack of competition. During the hot season work is 
carried*out under trying conditions; the least exertion causes 
profuse perspiration, the degree of discomfort depending mdstly 
on the nature of the work and on the surroundings, but is well 
pronounced, even in people who follow a sedentary occupation. In 
addition, during the very hot months of the year the continuous 
pouring out of perspiration during the day is not relieved by a 
respite during the hot night, when, although at rest with doors and 
windows wide open, one perspires freely and wakens up in the 
morning unrefreshed and more tired than one felt on going to bed. 



410 


It requires but little thought to realise that this discomfort plays 
a still more important part in the life of the women. Their work— 
domestic duties—is carried out during the hottest part of the day, 
indoors in the hottest part of the house, and in most instances in 
that part which has been most neglected in construction. It is very 
difficult to obtain domestic help, on account of the small number of 
domestic servants available, whose wages are in consequence very 
high, demanding an expenditure which is in most instances 
prohibitive for a medium income. The wife of the settler on the 
land is in a still worse plight, as she has not only to do her own 
housework, but in many instances has to cook for the employees. 
There is, in fact, no eight-hour working day for these women. 

As man is largely affected by his surroundings, and the quality 
and quantity of his work influenced by outer conditions, it can be 
readily understood that a tropical climate with a hot atmosphere 
laden with moisture, where the discomfort following any bodily 
exertion is great, is not conducive to a maximum output of energy, 
especially if the lack of competition makes the condition of life easy. 
There is no doubt that climate -per se tends to affect greatly the 
quality and quantity of physical and mental work. A cold climate 
conduces to physical activity, a warm climate to lassitude, and the 
economic conditions are then the determining factor for the output 
of energy. 

In North Queensland, with its remarkable natural resources, its 
remarkable recuperating power after a series of bad seasons, and its 
small population, the conditions of life are such that poverty is 
unknown and an easy living earned at a minimum cost of exertion, 
and the rate of wages is very high. 

Another factor which plays a very important part in the deter¬ 
mination of the quality of work is the abuse of alcohol, which is 
rampant amongst all classes of the community. A number of 
factors contribute to this abuse, such as the thirst caused by the 
heat, the temptation to quench this thirst owing to the great number 
of licensed public houses, the temporary stimulating effect of the 
slow poison, and last, but not least, the well-known open-hearted 
hospitality of the north, which finds its expression in 1 shouting.* 

The foregoing observations prove that in North Queensland both 
climate and economic conditions, namely, the lack of competition 



41 1 


and high wages, have created a tendency in the same direction, 
towards a deterioration of labour, and it is impossible at present to 
apportion correctly the influence of either factor. It is quite 
probable that, in the future, when the population of Northern 
Australia has increased and competition has become keener, the 
quality of labour may improve, unless in the meanwhile a definite 
decay of the race, due to climate, has set in. Up to the present, 
however, no such decay is apparent, but it must be kept in mind 
that in a newly-settled country, with a floating population, which 
to a great extent is maintained by immigration, racial degeneration 
would not make itself felt until after a somewhat prolonged period. 

The ‘Great Experiment of White Australia' in most previous 
discussions has been considered entirely from one point of view, 
namely, the health point. It has been assumed that the possibility 
of a population of European descent to live, propagate and thrive 
in a tropical climate, alone would decide the question. In Northern 
Australia, however, the economic conditions are of equal importance 
and present a problem as far-reaching in its influence as health for 
the permanent settlement of tropical Australia. 

LITERATURE 

Abbi Cleveland, quoted by Hann (1908). Vol. I, p. 57. 

Abon (190^ ). Philippine Journal of Science , B., Vol. IV, p. 225. 

-(1911). Philippine Journal of Science , B., Vol. VI, p. 102. 

Benedict (1915). Journal of Biol. Cb*m ., Vol. XX, p. 263. 

Blagden, and Fohdyce (i 775). Phil, Trans. London , Vol. XLV, p. 1, quoted from Schafer’s 
Text-book of Physiology, Vol. I, 1898. 

Breinl, and Priestley (1914). Annals of Trop. Medicine and Parasit . Vol. VIII, p. 606. 

Bruce (1916). Moisture and Temperature in Air in Relation to Comfort and Health. Reprint 
of paper read before the Public Health Section of the Roy. Soc., N.S.W., November 14, 
1916. 

Campbell (1917)* Journal Straits Branch , Roy. Asia. Soc. No. 76 

Chamberlain (191 x). Philippine Journal Med. Science. Vol. VI, No. 6. 

-and Vedder (191 k). Philippine Journal Science. B., Vol. VI, p. 403. 

Climes ha (1912). The Bacteriology of Surface Waters in the Tropics, London. 

Cuthbert (1911)- Journal Roy. Army Med. Corps , Vol. XVII, p. 56. 

Davy, John (1839). Researches , London. Vol. I, p. 161. 

-(1850). Phil. Trans., p. 437. 

Eijkman (1891). Virchow's Arcbiv. VoL CXXVI, p. 113. 

-(*893)* Virchow's Arcbiv. Vol. CXXXIII, p. 105. 

-(1895). Virchow's Arcbiv. Vol. CXL, p. 155. 

-(1896). Virchow's Arcbiv . VoL CXLIII, p. 448. 

Falks, L. (1907). Tropical Neurasthenia. American Journ. Med. Sci., April. 

Freer (1912). Philippine Journal Science , B. Vol VII, p. 2. 



Purnell (1878). Lancet. Vol. II, p. no. 

Gibbs (1912). Philippine Journal of Science, B. Vol. VII, p. 92. 

Glogner (1892). Virchow's Arcbiv. Vol. CXXVIII, pp. 160, 180. 

—— (1909). Arcbivfiir Scbiffs and Tropenbyg. Vol. XIII, p. 169. 

Haldane (1905.) Journal of Hygiene. Vol. V, p. 494. 

Hamm (1908). Handbucb der Klimatologie, Stuttgart. Vol. I, p. 57. 

Harrington, quoted by Harm (1908). Vol I, p. 57. 

Havelock, Charles (1913). Trans. Soc. Trop. Med. Hyg. Vol. VII, No. 1, p. 1. 

Hermann (1883). Arcbiv.f. Hyg. Vol. I, p. 1. 

Heyman, Paul, and Ercklentz (1905). Zeitscbrift fur Hyg. Vol. XLIX. 

Hill, L. (1914). Reports Local Government Board, No. 100. New Series, London, p. 18. 
-(1916). Pbil. Trans., p. 214. 

-and Flack (1909). Proceedings, Phys. Soc., Jottrn. of Physiol. VoL XXXVIII. 

Hill, Flack, Rowlands, and Walker’ (1913). Smithonian Miscellaneous Collections. Vol. 
XL (quoted by Hill, 1914). 

Hunt, Atlee (1915). The Northern Territory of Australia. Memorandum. Melbourne. 
Joussrr (1884). Traitk de Vacclimatement et de l*acclimatation, Paris. 

McCay (1912)* The Protein Element in Nutrition, London, p. 156. 

Marestang (1889). Arch. de Mid. Navale, No. 2. 

Musorave, and Sison (1910). Philippine Journ. Science , B. Vol. V, p. 325. 

Neuhaus (1893). Virchow's Arcbiv. Vol. CXXXIV, p. 365. 

Osborne (1910). Journal of Physiol. Vol. XLI, p. 353/ 

•-(1916). Proc. Roy. Soc. Viet., N.S. Vol. XXIX. 

Pembrey (1898). Animal Heat. Schifer’s Text-hook of Physiology. Vol. I, p. 812. 

-(1913). Journal Royal Army Med. Corps. Vol. XXI, p. 156. 

Phalen (1910). Philippine Journ. of Science, B. Vol. V, p. 525. 

Plehn (1892). Virchow's Arcbiv. Vol. CXXIX, p. 285. 

-(1898). Die Kamerun Kftste. 

-(1906). Arcb.f. Scbiffs und Tropenbyg.V ol. X, p. 220. 

Price, Basil (1913). British Med. Journal, November 15, p. 1290. 

Ranke (1900). Quoted by Schilling (1909). 

Rattray (1870.) Proc. Roy. Soc. Vol. IT, p. 513. 

Sambon (1907). Journ. Tropical Medicine and Hyg. Vol. X, p. 67. 

Van der Scheer (1890}. Geneesk. Tijdscbr. v. Nederl. Ind. VoL XXX. 

Schilling (1909). Arcbiv. f. Scbtffs und Tropenbyg. Vol. XIII, p. 1. 

- (1909). Tropenhygiene, Leipzig, p. 514. 

Schmidt (1909). Arcbiv. fur Hyg. Vol. LXIX. 

——— (1909). Quoted by Aron (1911). p. 106. 

Shaklee (1917). Philippine Journal Science, B. Vol XII, p. 1. 

Sutton, Harvey (1908). Journal Path, and Bact. VoL XIII, p. 68. 

Taylor, Griffith (1916). The Control of Settlement by Humidity and Temperatures. 

Commonwealth Bureau of Meteorology. Bulletin 14. 

Thornley (1878). Lancet. VoL I, p. 554. 

Wick (1910). Arcbiv. f. Scbiffs und Tropenbyg. Vol. XIV, p. 106. ^ 

Wickline (1908). MHit. Surgeon. Vol. XXIII, p. 283. 

Woodruff (1905). The Effects of Tropical Sunlight upon the White Man, New York. 
Young (1915). Journal of Physiol., Vol. XLIX, p. 222. 

-(1919). Composition of Urine in Tropical Australia. Ann. Trop. Med. & Parasitol 

Vol. XIII, p. 215. 

Zuntz, and Tendlat, quoted by Pembrey (1913^. 



4*3 


THE EXPERIMENTAL INFECTION IN 
ENGLAND OF 

ANOPHELES PLUMBEUS, Stephens, 
AND ANOPHELES BIFURCATUS, L„ 
WITH PLASMODIUM FIFAX 


BY 

B. BLACKLOCK 

AND 

HENRY F. CARTER 
(Received for publication io February , 1920) 

HISTORICAL 

The potentiality of A. plumbeus as a malaria carrier has 
been variously assessed. Theobald (1901)/ in his Monograph of 
Culicidae, remarks that both A . plumbeus (= A. nigripes) and 
A. bifur cat us are malarial carriers, but are not nearly so abundant 
as A . macultpennis in England. Nuttall and Shipley (1901) give 
‘A. nigripes ? 9 as a carrier of malaria. Bacot (1918) says that 
there is no evidence that this species is able to carry malaria. Grove 
(1919), in a report to the Local Government Board, states 
‘A. plumbeus from the rarity of its occurrence is certainly of very 
little danger. It has not yet been proved whether it is capable of 
transmitting the malarial parasites.’ 

These apparently irreconcilable statements confronted us in the 
commencement of our investigation, and it became necessary to 
ascertain which view should be considered the correct one. On 
making a search through the literature we found no mention of 
experimental work associated with A . plumbeus (vel ni gripes) 
beyond that in the early Italian records, more especially the works 
of Bastianelli, Bignami and Grassi. These workers (1898) dissected 
amongst other mosquitoes one specimen which they identified as 
A . nigripes; the mosquitoes were caught in the room of four patients 

* For literature consulted, see pp. 441-444. 


suffering from malaria, probably malignant tertian; the specimen of 
A. nigripes was not infected. Grassi, Bignami and Bastianelli 
(1899 2 ), in giving an account on February 5th of their work during 
the month of January, 1899, say that the need for experimenting 
with A. bifurcatus and nigripes has become evident, and that they 
propose to carry out such experiments within a short time. This 
work was evidently accomplished by June 18th of 1899, because on 
that date Grassi (1899) felt himself justified in stating that it was 
then proved that all the Italian species of the genus Anopheles 
propagate malaria. 

It is between February 5th and June 18th, 1899, therefore, that 
we may expect to find the evidence upon which Grassi made this 
somewhat comprehensive statement, and the following is the evidence 
which we have obtained. Grassi, Bignami and Bastianelli (1899 1 ) 
wrote, in May:—‘We have together made experiments with 
Anopheles bifur cat us var. ni gripes with blood containing gametes 
of tertian fever and with blood containing crescents; the results were 
undoubtedly positive.’ 

They amplify this bald statement in another place (1899 3 ) on 
May 7th as follows : —‘ We have completed a series of experiments 
on A. bifur catus (1). The individuals which were used inclined to 
the variety nigripes. We have experimented with these insects on 
a case of crescent infection and on cases of tertian infection. Both 
with crescent and tertian infections we have obtained positive 
results. The various stages of development observed in the walls 
of the intestine of A. bifur catus correspond perfectly with those 
already noted by us in A. claviger , kept under the same conditions 
of temperature and examined at the same interval after the time of 
biting.’ 

These statements by Grassi and his collaborators are the sole 
evidence which we have been able to obtain of any reference to 
A. plum bens (vel nigripes') in experimental work on malaria, and 
it appears to us, apart altogether from the unfortunate lack of detail 
in the description of the experiments themselves, that the views of 
the authors upon the identity of the mosquito with which they were 
working are of great interest. It will be noted that a change in the 
nomenclature of the mosquito has occurred during this short period. 
First of all there are references to A. nigripes , next comes a reference 



4i5 


to A. bifurcatus var. nigripes, and in the conclusion of the account 
of the above experiment the mosquito is referred to simply as 
* 4 . bifurcatus. The attitude of the authors towards the question of 
species is expressed by them (1899 1 ) thus:—‘Comparative observa¬ 
tions made by Grassi in Calabria had led 'him to suspect also the 
other species of Anopheles found in Italy of being agents in 
transmitting malaria. These species, according to the researches 
of Ficalbi completed by Grassi, are the following: — 

(1) Anopheles bifurcatus (there is a variety called A. bifurcatus 

var. nigripes ). 

(2) Anopheles pseudopictus (Grassi). 

(3) Anopheles superpictus (Grassi).’ 

From this it would appear that they did not recognise 
A. nigripes as a separate and well defined species, but only as one 
particular form of A. bifurcatus. That this is in reality their 
considered opinion is made more evident by an important footnote 
by Grassi, indicated above (1), which runs:—‘After the examina¬ 
tion of a certain number of individuals collected in different 
localities, I hold with Ficalbi that A. nigripes is simply a variety of 
A. bifurcatus , an opinion which I intend to confirm by the examina¬ 
tion of numerous individuals as well as of the larvae and ova.’ No 
modification of this opinion on the part of Grassi has been traced 
by us. 

From a study of the foregoing facts, therefore, we gain the 
information that: — 

(1) This series of experiments of Grassi and his collaborators 
was carried out with wild mosquitoes, and not laboratory-bred 
insects. 

(2) Whatever the true identity of the mosquitoes which they were 
using in these experiments, infection of the gut only was obtained. 

(3) Grassi and his co-workers almost certainly experimented, not 
with A. plumbeus —a tree-hole breeder, but with a dark variety of 
A. bifurcatus. 

Grassi, moreover, was aware that individuals of A. bifurcatus 
varied considerably in appearance, because he states explicitly 
(i 9 ° 3 ) that in June, 1899, he had, by himself, experimented 
successfully with ‘ tAnopheles bifurcatus tipico.' 

It is of interest to note that at the time when the experiments 



4i 6 


recorded above were done by Grassi, Bignami and Bastianelli, 
Grassi had not observed mosquitoes breeding in tree-holes. Not 
until 1901 do we find any reference by him to such breeding-places, 
and when he did come upon them he referred to them very shortly 
under the heading A. bifurcatus, ‘Ich fand in einer mit Wasser 
angefiillten Ausholung eines Baumes, larven des A. bifurcatus var. 
nigripes' He shews in this statement no appreciation of the fact 
that this tree-hole breeder was a separate and definite species, nor 
that the adults and larvae of it were quite distinct from those of any 
known A. bifurcatus , whether small and dark, or large and ‘tipico.’ 
Not only is it evident that Grassi failed to recognise the importance 
of his capture, but in our opinion this was the only occasion upon 
which he encountered the true A. ni gripes (i.e., plumbeus). 

Galli-Valerio and de Jongh (1903) did not associate this observa¬ 
tion of Grassi’s with the existence of a different species, because they 
say: —‘ We have often obtained from larvae very small forms of 
A. bifurcatus presenting the characters of A. ni gripes, Staeger, but 
having observed all the range of variation in size between typical 
A. bifurcatus and typical A. ni gripes, we agree with the opinion of 
Ficalbi, who says that it is not a case of a distinct species, but simply 
a form of Anopheles bifurcatus, which is liable to present very great 
variations in size.’ Later, however, Galli-Valerio and de Jongh 
(1913), discussing their previous identification of Anopheline larvae 
in tree-holes in Plamont as * 4 . bifurcatus, state that the tree-hole 
breeding larvae are A. ni gripes-. —‘This is the first time,’ they say, 
‘that the species has been found in the Canton of Vaud in 
Switzerland, and what we have previously called A. bifurcatus var. 
ni gripes is simply a small form of bifurcatus.' They refer to the fact 
that Blanchard also was under the impression that A. bifurcatus 
and A. ni gripes were identical. From this time onwards Galli- 
Valerio has studied A. plumbeus with great care, and yet recently 
(1917 2 ) he writes, ‘Mes recherches precedentes ont demontre 
l’existence en Suisse de trois especes d’Anophelines: A. maculi- 
pennis, A. bifurcatus et A. ni gripes. Tandis que pour les deux 
premieres especes, leur role dans la transmission de la malaria est 
demontre, la troisieme n’a pas encore fait l’objet de recherches 
exp£rimentales a ce point de vue; mais il parait probable qu’elle 
intervient aussi dans la contamination et dans la transmission 
malarique, surtout dans les zones boisees.’ 



4*7 


As a result of our examination of the literature on this subject, 
we agree with the opinion of those who up to 1919 held that 
A. plumbeus had not been convicted by any evidence, experimental 
or other, of being an agent in the carriage of malaria. 


EXPERIMENTAL 

i. Experiments with laboratory-bred Anopheles plumbeus 

In a preliminary note (1919) we recorded the experimental 
infection of the gut and salivary glands of Anopheles plumbeus bred 
in the laboratory from pupae collected in England, and fed upon 
a patient whose blood contained Plasmodium vivax . More detailed 
information with regard to those experiments and additional ones 
conducted along the same lines are given in the present paper. 

The work was carried out at the Liverpool School of Tropical 
Medicine during the period July to October, 1919. No experi¬ 
ments on the transmission of the disease to healthy human beings 
were made, because transmission experiments appear almost 
superfluous when such work as that of Mitzmain (1916) is taken into 
consideration. In his work with Anopheles punctipennis , Mitzmain 
found that a single mosquito which had sporozoites in its salivary 
glands was capable of infecting a series of healthy persons by single 
bites, and this even when the insect did not have an opportunity of 
taking a full meal of blood in biting. 

Technique . Breeding . In the case of the first experiments the 
larvae of the pupae were obtained from Grasmere in the Lake 
District, they were taken from the tree-hole in which they were living 
and placed in a jar together with a quantity of the water and debris 
present in the cavity in the tree trunk, and brought to the laboratory 
by motor car; they withstood the journey well, and a number of 
adults subsequently emerged. An attempt made, at a later period, 
to send live pupae on moist moss from Wales by post proved 
unsuccessful; all the pupae died on the way. Larvae were found 
to be capable of surviving days of travelling if placed in bottles or 
tubes half-filled with the water from the tree-hole in which they 
were found. Our experience so far points to the greater hardiness 
of young larvae in this respect. When the material was brought to 
the laboratory it was placed in suitable glass breeding vessels and 



covered over with bell jars into which the emerged adults could 
ascend; in order to approach as closely as possible the natural 
conditions, the bell jar was encased in brown paper. Transference 
from the bell jar to the feeding apparatus was effected as required 
by means of a piece of wide glass tubing, over one end of which 
netting was fixed; the bell jar having been removed under mosquito 
netting, the mosquitoes, one at a time, were captured in the tube 
and then liberated into the feeding apparatus, which consisted of 
a wide glass cylinder covered at each end by netting kept in position 
by rubber bands. The cylinder was set upright in the bottom of 
a Petri dish, on which a piece of moist blotting paper was placed. 
The food provided, apart from the single meal of infected blood, 
consisted of raisins. Each day the blotting paper was moistened 
and a split and moistened raisin placed on the netting on the top 
of the feeding jar. The netting on the feeding apparatus and the 
blotting paper in the Petri dish were renewed daily at first, but 
later about every two days was found sufficient. In the incubator 
experiments a vessel containing water was placed in the incubator, 
and as evaporation proceeded fresh water was added. 

Feeding. Anopheles plumbeus rarely attempted to feed on 
blood within the first twenty-four hours of emergence from the pupa, 
but after that period most of them fed readily and engorged them¬ 
selves with blood, some even feeding well when it was obvious that 
they had recently partaken freely of the raisin diet provided. 

Dissection. All mosquitoes which died after the infected feed 
were dissected, and at intervals others were killed and examined; 
fresh preparations of the mid-gut and salivary glands were made 
and examined in 0*85 per cent, sodium chloride; for permanent 
preparations the gut and glands were fixed in Bless* fluid, stained 
with haematein and differentiated in acid alcohol. 

The results obtained by feeding and dissection are given below 
in Table I. Gametes were present in the peripheral blood of all 
patients used for the feeding experiments. The case upon which 
Experiments A and C were done had at the time of feeding 
numerous parasites in the peripheral blood and had a temperature of 
103° F. 

It is worthy of note that salivary gland infection was present in four 
of eight mosquitoes which were dissected after ten days or longer at 28° C. 
from the time of the infected feed. 



+I 9 


Table I. Showing the mult* of feeding laboratory-bred A. plumbeus 
on patients infected with P. vivax. 



Date 

fed 



Number 



Temperature 

used 

Remarks 

Exp. 

Fed 

Dissected 

1 

Infected 

Gut 

Sal. 

Glands 

A ’ 

11.7.19 

2 

2 

0 

0 

0 

Room 

One, allowed to live, survived 71 
days after infected feed 

B. 

10.7.19 

6 

6 

3 

3* 

2 

28 °C. 

C. 


2 

2 

0 

0 

0 

28 °C. 


D. 

11.7.19 

2 

2 

1 

1 

0 

Room 


E. 

* 4 - 7 * *9 

l 

6 

5 

3 

3 

; 

2 

z8 a c. 

One died 1.8.19 but could not be 
dissected. 


* One of these, which had not yet infection of the salivary glands, was killed on the 8th day. 
In four of the twelve oocysts found in the gut, sporozoites were observed. 


2. Experiments with laboratory-bred Anopheles bifurcalus 

In order to obtain some idea of the relative infectibility of 
A. plutnbeus and one of the better known anophelines which occur 
in England, we made some experiments with A. bifurcatus under 
the conditions which obtained in procuring salivary gland infection 
in A. plumbeus , with the exception that the experiments were carried 
out at a later period of the year and that the same patients were not 
used. The results are given in Table II, and are of interest. 
Experiment Q, in this control series, is especially interesting, 
because in this case the maturity of the gametes was tested by 
observing exflagellation, which was found to be proceeding freely 
before feeding was carried out. 


Table II. Showing the results of feeding laboratory-bred A. bifurcates 
on patients infected with P. vivax. 


Exp. 

Date 

fed 

Number 


Temperature 

used 

Remarks 

Fed 

Dissected 

Infected 

Gut 

Sal. 

Glands 

M. 

23.9.19 

2 

2 

0 

o 

0 

28V 


N. 

29.9.19 

8 

8 

c 

0 1 

o 

28 °C. 


O. 

17.10.19 

12 

12 

1 

1 

0 

1 1 

28 °C. 




420 


In this experiment, thirteen mosquitoes were dissected, after 
ten days or longer at 28° C. from the time of the infected feed; no 
instance of salivary gland infection was obtained. 

We hope to have an opportunity in the summer of making a 
rigidly comparative series of experiments with A. plutnbeus and 
A. bifur catus in order to determine with more accuracy their ability 
to acquire infection in laboratory experiments. 

4 Previous experimental work with these species 

The amount of such work recorded in the literature is very small. 
Neither of the two species used by us has previously been experi¬ 
mentally infected in this country, nor has either of them ever been 
found infected in nature. With regard to experiments conducted 
on mosquito infection in England, the only record is that of James 
(1917) who worked with Anopheles maculipennis. The mosquitoes 
which he used were apparently adults captured in huts and bed¬ 
rooms of malaria patients at Sheemess; they were fed on a benign- 
tertian gamete carrier, and afterwards dissected. Infection was 
obtained in two mosquitoes, and was in each case confined to 
the gut. 


SUMMARY 

We have been able to infect laboratory-bred A. plutnbeus with 
P. vivax. At 28° C. infections of the gut and salivary glands were 
obtained; at room temperature (max. 26° C., min. I7°C.) gut 
infection only was obtained. We have also produced infection of 
the gut with P. vivax in the case of A . bifur catus at 28° C. 

This is, we consider, the first experimental evidence produced 
that A . plutnbeus is capable of becoming infected with a malaria 
parasite. As regards A . bifur catus we have proved that in England 
the native form of this mosquito is capable of being infected with 
malaria at 28° C. 



4 11 


OBSERVATIONS ON ANOPHELES 
(COELODIAZESIS) PL UMBE LW, Stephens, 
WITH SPECIAL REFERENCE TO ITS 
BREEDING-PLACES, OCCURRENCE IN 
THE LIVERPOOL DISTRICT, AND 
POSSIBLE CONNECTION WITH THE 
SPREAD OF MALARIA 


BY » 

B. BLACKLOCK 

AND 

HENRY F. CARTER 
(Received for publication 16 February , 1920) 

Plates X— XII 

As will be gathered from the preceding paper, we have, during the 
last few months, given considerable attention to Anopheles plumbeus, 
Stephens, the bionomics of which species are probably less known 
than those of any European Anopheline mosquito. This lack of 
knowledge is aggravated by the scattered nature of the literature 
pertaining to this insect and by the not infrequent contradictory 
statements made regarding it. In recording our observations, 
therefore, it will not be inexpedient if, besides alluding to the more 
important work performed by other observers, we give some general 
account of the species. 

Synonymy 

Anopheles plumbeus was discovered by Haliday in the north 
of Ireland in 1828. But although this author gave a brief 
description of his specimens, he evidently doubted their being 
distinct from . 4 . bifurcatus, L., and did not name them. Stephens, 
however, in the same journal and in a paper immediately following 
Haliday’s assigned the name plumbeus to the description supplied 
by the latter writer. 



422 


In Denmark, in 1839, Staeger captured a female Anopheline 
which he described as A. nigripes. But the apparent rarity of 
A. nigripes and the resemblance of the adult to small dark forms of 
A. bifurcatus led to much confusion, and, until Eysdl (1912) 
described its early stages and Galli-Valerio and de Jongh (1912 2 ) 
its breeding habits, most continental authorities considered it a 
variety of this species. Such a conclusion was based chiefly upon 
incorrect identifications. Theobald (1901), however, not only 
recognised its validity but associated it with Stephens’ species, a 
suggestion which Edwards (1912) accepted and extended by 
resuscitating the name A. plumbeus. 

James* (1911) described A. barianensis from the Western 
Himalayas. But Christophers (1916), after studying all its stages, 
decided that it was identical with the European form of 
A. plumbeus. This author also suggested that the North American 
A. barberi, Coq., would probably prove synonymous with 
A. plumbeus , but in the opinion of one of us (H. F. C.) the species 
are quite distinct, as the descriptions and figures of the American 
species given by Smith (1906) and Howard, Dyar and Knab (1912- 
1917) reveal important morphological differences. 

The synonymy of A. plumbeus is therefore: — 

Anopheles plumbeus , Stephens (1828) 

Anopheles tu gripes, Staeger (1839) 

Anopheles barianensis, James (1911) 
and to that of A. bifurcatus must be added : — 

Anopheles nigripes, Ficalbi et auct. (nec Staeger) 

Anopheles bifurcatus var. nigripes, Ficalbi et auct. (nec 
Staeger). 

Relationship with other Anophelines 

A. plumbeus, Steph., A. barberi, Coq., and the south Indian 
A. culiciformis, Cogill, compose a group of species all of which 
show great selectivness in their habits.* They are primitive forms, 
and thus are related to the ‘Anopheles’ (sens, str.), 1 Stethomyia’ 
and ‘ Myzorhynchus ’ groups. Their eggs and larvae are peculiar, 
and present characters which have been considered of generic or 
sub-generic value. Dyar and Knab (1906) founded the genus 
Coelodiazesis on the larval characters of A. barberi, but subsequently 

* See note on p. 444. 



4*3 


Dyar (1918) reduced it to sub-generic rank. Independently, Eysell 
(1912) created the genus Cyclophorus for A. plumbeus on account 
of the striking characters of its eggs. 

* 

General Distribution 

From what has been said above, it will be seen that A. plumbeus 
is very widely distributed. Owing to its selective habits and our 
comparatively recent discovery of them, records are meagre and, 
for the most part, disconnected. But it has been found in nearly 
all the European countries, including Scandinavia and Italy, and 
also in the Western Himalayas. The records show that it occurs at 
varying altitudes up to 8,000 ft., and apparently that it is restricted, 
or almost restricted, to forested regions. Yet it has been captured 
in towns, e.g., Edinburgh, Grenoble, and in the suburbs of London, 
Paris and Bonn. Its supposed rarity has been due to the paucity 
of adults collected or observed, but since the discovery of its 
breeding-places it is becoming evident that it is much more common 
than was believed. 

Characters of the Adult and Immature Stages of A. plumbeus 

The adult has been described by several systematists since 
Haliday’s time, and the early stages in some detail by Eysell 
(1912), Galli-Valerio and de Jongh (1912 2 ), Martini (1915), 
Christophers (1916), and others (cf. p. 432 re Meinert’s description). 
It will, therefore, be necessary to mention only the more important 
diagnostic characters. 

A. plumbeus measures from 4 mm. to 5 mm. in length and is of a 
general dark colour with unspotted and densely scaled wings. The 
palpi are uniformly dark in both sexes, and the hair-tuft on the 
last two segments in the male is much reduced. The thorax, as in 
A. barbcri and A. culictformis, is relatively shorter in comparison 
with the width (not more than twice as long as wide) and more 
rounded than in other Anophelines; the median region of the 
mesonotum is broadly ashy-grey, with a central line of white scales 
anteriorly, and is sharply separated from the sides which are a 
rich dark brown colour. The legs are very dark, with inconspicuous 
pale spots at the apices of the femora and tibiae. The abdomen is 
blackish brown and hairy. 



4 2 4 


The egg measures o*5-o*6 mm. long by o*2 mm. wide. It is 
fusiform, broad in the middle and tapering sharply towards the 
^ends, and is completely surrounded by a float of nearly uniform 
width. 

The larva is dark, almost black, above, and whitish below, and 
when mature measures 5 mm. in length. As with the other members 
of this group, it is characterised by possessing long branched hairs 
on abdominal segments 1-6 inclusive, and by the presence of minute 
hairs only on the head. The clypeal hairs are simple, the antennae 
are almost devoid of minute spines on the shaft and the basal hair 
is very small. The abdomen bears dorsally five pairs of palmate 
hairs and laterally on segments 1-7 conspicuous radiate bristles, 
each composed of from three to seven setae. The ninth segment is 
strongly chitinised. 

The pupa, like the larva, is pale ventrally; but unlike it, shows 
poor development of the abdominal hairs and spines. 

Adult Habits 

Comparatively little is known of the habits of the adult 
A. plumbeus. It is generally considered to be an essentially sylvan 
and 4 wild * species, which occurs but rarely in houses. In woods, 
however, when observed at all, it has been recognised as a vicious 
and persistent biter, and Edwards (1916) states that it is a 
4 common blood-sucker in wooded districts in this country, often 
biting during the day/ For the most part, the statements made 
regarding the slight tendency of this mosquito to invade houses are 
based upon the occasional capture of one or two individuals in such 
places, but from the few serious observations made it would seem 
that this species is in no way averse to entering houses. Thus 
Burton (see Lang (1918)) states that near Shrewsbury A. plumbeus 
is not rare, 4 It bites all day and night; I have never seen more than 
two trying to bite at the same time. It is found mostly in the woods, 
but will come into houses to bite more often than will A. bifurcatus .* 
Hesse (1918) says that the adults enter houses voluntarily. 

That A. plumbeus has not been recorded more frequently in 
houses is, in our opinion, due chiefly to the fact, noted by 
Christophers (1916), that the females retire to the shelter* of their 
breeding-places shortly after feeding. Unlike ,A . maculipennis , 



4*5 


therefore, they will remain but a short time in the house, and the 
chances of seeing them except in the mornings and evenings will be 
small. In this connection Christophers, working with A. plumbeus 
in the Western Himalayas, states: ‘ The adults may be found during 
the day resting inside hollow trees. Most of the females taken in 
such situations contained blood in the gut, and as it was ascertained 
that in the mornings and evenings, but more especially the former, 
the female entered houses and bit freely, it is probable that such 
blood was to a large extent of human origin.’ It is evident, 
therefore, that in this country much closer observation of the feeding 
habits of A. plumbeus is necessary. 

A. plumbeus bites readily in the laboratory twenty-four hours 
after emergence, and will do so at any time of the day and in full 
daylight. Females, kept in glass cylinders, became engorged in 
from one and a half to four and a half minutes, but three minutes 
was the usual period of time required. Their avidity for blood was 
shown in several instances by females, which had distended them¬ 
selves with blood half an hour to one hour earlier, re-engorging for 
a period of one minute or so. The bite was singularly devoid of pain 
and produced very slight local reaction; the usual character of the 
local reaction was the appearance of small white circular areas, each 
surrounded by a reddened zone, and this appearance was noticed 
within an hour, or up to ten hours, after the bite. The bite of a 
mosquito and its effect is largely a question of individuality, but it 
is worthy of note that, with some people at least, the bite of 
A. plumbeus may pass entirely unnoticed, and for a supposedly 
‘ wild ’ species this is of interest. In our experience the bites of 
A. bifurcatus and the purely sylvan O. nemorosus or 0 . maculatus 
are distinctly more painful. 

The great majority of the females under observation were not 
kept for the purpose of studying their biology and were only given 
one meal of blood each and often killed when perfectly vigorous. 
They fed readily on moistened raisins, however, and in several 
instances lived considerable periods on such diet. 

Of twenty-four females which received one meal of blood and 
had free access to raisins: five lived less than ten days (one being 
killed); eleven lived 10-20 days (six killed); two lived 20-30 days 
(one killed on the 28th day); four lived 30-40 days (one killed on 



4 26 

the 40th day); and two lived over 40 days. Of the last two, one, 
although active and vigorous, was killed (Mr the 48th day, and the 
other died on the 75th day.* Most of the females had been kept in 
an incubator at 28° C., but the last mentioned had remained in the 
room, the temperature of which varied during its life from I2°C. 
to 26° C. 

Of three females which received raisin diet only and were kept 
at room temperature (17 0 C.—2i°C.) one lived 14 days and two 
15 days each. 

Of nine raisin-fed males the lengths of whose lives were ascer¬ 
tained, five died in less than 10 days, and the remaining four lived 
29, 3 1 < 33 > and 47 days respectively. These four were all confined in 
cylinders with one or two females, and during their lives the last 
two were subjected to temperatures ranging from 12° C. to 28° C. 

Mr. R. F. L. Burton of Shrewsbury has made many interesting 
observations on the swarming of the males of A. plumbeus. He 
states (in Lang (1918)) that on windless evenings from July to 
October they may always be found swarming in certain definite 
spots, and, in a personal communication, informs us that one such 
spot is about 100 yards from an elm-stump, the only breeding-place 
known to him. He states that this swarm usually consists of seven 
or less males and that it occurs on the outskirts of a wood and six 
feet from a slow, muddy stream. Mr. Burton has observed in other 
spots swarms consisting of as many as thirty individuals. 

Breeding-places 

A . plumbeus and the other species of Coelodiaeesis are essentially 
tree-hole breeders, their larvae developing in the water contained 
in rot-holes. Many authorities—Martini (1915), Galli-Valerio 
(1917 2 ), Eckstein (1918 1 )—agree that they live exclusively in such 
places, and, unless situations be found where the biological condi¬ 
tions approximate to those peculiar to the rot-holes, they are 
probably correct. Apart from records which undoubtedly pertain 
to dark forms of A. bifurcatus, we have found three relating to the 
discovery of A. plumbeus larvae in places other than rot-holes. 
Evans (in Theobald (1910)) claims to have bred males from larvae 


* This mosquito has been referred to in Table I of the preceding paper. 



taken fiom a pool near Lasswade, Edinburgh, but as this gentleman 
made no reference to the striking characters of the then unknown 
larva, this record is not generally accepted. C. Blanchard (1918) 
described a young larva, which he suggested was that of 
A. plumbeus (nigripes), from a marsh, but later R. Blanchard (1918) 
stated that an error of identification had been made and that 
the specimens captured were the first stage larvae of A . maculipennis. 
The third record is deserving of much closer attention and was made 
by Theobald (1908). This author writes: ‘ We have found such 
as A. nigripes, Staeg. and A. bifurcatus, L. breeding in the water 
of peat cuttings in Wales and Somerset.’ He has recently informed 
us privately that his identifications were made from adults reared 
from the larvae found. In an attempt to confirm this observation 
we recently examined the water of peat cuttings in this neighbour¬ 
hood but without success, although in a hole in a birch tree, situated 
not more than 15 yards from the peat water, hibernating larvae of 
A. plumbeus were discovered. In this case, however, the water in 
the cutting was almost colourless and very different in appearance 
from that in which A. plumbeus larvae are usually found; but if, as 
is conceivable, the water in very old cuttings acquires a greater 
resemblance to that of the rot-holes, the possibility of its acting as 
a breeding-place must be considered. 

A very good idea of the type of breeding-place selected by this 
Anopheline may be gained from Plates X and XI and from the table 
on pages 430 and 431. We have on one occasion only found the larvae at 
ground level (cf. Plate XI, figs. 3 and 4), and then the breeding-hole 
was well sheltered in the projecting portion of a root and by a mass 
of fallen leaves which almost completely covered it. The character¬ 
istic hollows formed by the root-buttresses of beech trees containing 
collections of clear water of a temporary nature have never, yet 
yielded us A. plumbeus larvae; but in one instance an exceptionally 
deep hollow contained, so far as we could ascertain, four larvae of 
O. geniculatus. Galli-Valerio (1916) states that the larvae may also 
occur in holes in stumps of trees and in the forks between the 
boughs of beeches. In most cases examined by us the breeding- 
place has provided ample shelter for both larvae and adults, even 
although the tree itself was in an exposed position. In several 
instances the orifice of the hole was almost blocked by a piece of 



428 

broken and rotten branch which came away.at the slightest pull; or 
in private grounds and parks where the trees were more carefully 

tended it was occasionally found that potential breeding-places were 
present in sawn-off branches which, a few yards away, appeared 
perfectly sound. On investigation, a small orifice was visible at 
the lower edge of the cut surface which had rotted away behind and 
below, leaving internally a fair-sized hole. The depth of some of 
the rot-holes is considerable, sometimes three feet or more, so that, 
bearing in mind the frequently admirable shelter provided and the 
resultant protection from evaporation, the permanent nature of most 
of these breeding-places is easily understood. Eysell (1912), who 
at the time did not know of the type of breeding-place selected by 
this insect, remarked that in the abnormally hot dry summer of 1911 
he was able to obtain more A. plumbeus material than ever; and 
he states that in this season only those mosquitoes breeding in 
permanent water reservoirs were numerous. Howard, Dyar and 
Knab (1917), speaking of Anopheles barberi, say ‘This species 
occurs, with other mosquitoes breeding in tree-holes, in forested 
regions, often when the country is so dry that no other mosquitoes 
are found.’ 

Christophers and Khazan Chand (1916) found the larvae of 
A. culiciformis in about 10 per cent, of the tree-holes holding water, 
and as a rule this water was of a ‘characteristic deep brown-tint 
like strong tea or coffee.’ We have made no estimate regarding the 
percentage of tree-holes harbouring A. plumbeus larvae, but the 
remark of these authors regarding the colour of the water is equally 
applicable to this species ; we have never yet found larvae in holes in 
which the water was colourless (on filtration). The exact colour 
of the water in which larvae occur appears to vary from yellowish- 
brown to deep reddish-brown, and is evidently a solution of sub¬ 
stances derived from the rotting tree and vegetable debris at the 
bottom.* Galli-Valerio (1916) states that in the laboratory the larvae 
develope better if kept in dark yellow containers and adds (1917 1 ) 
that the water contains tannin. The depth of the surface water in 

* An analysis of a sample of this water (specimen i) made by Professor W. Ramsden, 
of Liverpool University, is given on p. 440. Specimen 2 was obtained from what appeared 
to be a suitable breeding*place, but no larvae were found. The absence of larvae from such 
water was so exceptional that we requested Professor Ramsden to examine it also. 



the rot-holes varies greatly and may not be more than a few inches. 
Even in deep holes the accumulation of vegetable matter is sometimes 
so large that little water is evident. 

From the table on p. 430 it will be seen that breeding-places have 
been found in various kinds of trees; some of these have already 
been recorded by other observers as harbouring larvae. To the list 
must be added silver-fir (.Abies pectinata ) (Galli-Valerio and 
de Jongh (1915)) and hornbeam (Carpinus betulus) (Mr. F. W. 
Edwards, personal communication). Apparently, therefore, the 
species of tree is only of importance from its tendency to form 
suitable breeding-places. Certain kinds appear to show a higher 
proportion of rot-holes in the trunks while others, particularly when 
old, may by their conformation offer more opportunities for the 
retention of water. Beech trees have been especially referred to in 
connection with A . plumbeus and old beeches possibly are more 
liable to form breeding-places than other species of tree. In our 
investigations, and more particularly those conducted in the vicinity 
of Liverpool, the search for breeding-places was largely confined 
to comparatively small trees (the diameter of the main trqnk seldom 
being more than eighteen inches), and of these sycamores easily 
ranked first in providing us with larvae. Near Simla Christophers 
(1916) found that oaks were most favourable. 

The continual association, in the literature, of A. plumbeus with 
forests and woods is natural, in view of the facts now known 
concerning its habits, since in such places greater opportunities for 
breeding occur, and the chance of encountering adults is increased. 
But although this mosquito is always associated with trees it must 
not be identified solely with forests or woods. We have found 
larvae in trees situated in isolated spinneys, in meadows, or forming 
with two or three others a small clump—in situations, in fact, which 
could not by any stretch of imagination be termed woods. Further, 
they may and do occur in trees growing near habitations, both in 
rural districts and in towns, wherever suitable breeding-places exist. 

The difficulties attending the search for breeding-places of 
A. plumbeus are many and various. The work is arduous and 
frequently involves climbing and the use of special apparatus for 
sampling the water. Ladles and spoons of various sizes, bent at 
different angles, may be all that is required, but syphons are 



TABLE I. 

Details of the breeding-places observed. 


Locality 

Kind of Tree 

Height of 
aperture 
above ground 

Dimensions of 
rot-hole, in inches 
(approx.) 

Situation 

Remarks 

ENGLAND 
Liverpool District 
Hightown, Lancs. 

Beech 

{Fagus stlva tica) 

5 {L 

8 x 9 x 18 deep 

In small wood with 
dense vegetation ; 
200-303 yards from 
nearest houses. 

7.6.1919. Larvae of 
various sizes and 
pupae found, also 
many larvae and 
pupae of O. geni- 
culatus. 

26.7.1919. No larvae 
or pupae found. 

—.10.1919. No larvae 
or pupae found. 

Fazakerley, Lancs. ... 

Elm 

(Ulmus campestris) 

8 ft. 

19 x 12 x.17 deep 

In private grounds ; 

12 yards from 
entrance to house 

1.2.1920. Numerous 
small larvae of 

A. plumbeus found. 

Knowsley Moss 
(By kind permission 
of D. Hamilton, 
E^.) 

Birch 

{Betula alba) 

7 ft. 

6 x 6 x 12 deep 

200 yards from lodge 
and 10-15 yards 
from peat cutting 

7.2.1920. Many 
small larvae of 

A . plumbeus. No 
mosquito larvae 
found in water 01 
peat cutting. 

Roby. 

(By kind permission 
of D. Hamilton, 

£»q-) 

Sycamore 
{Acer pseudo - 
platanus) 

3 ft. 

14 x 8 x 12 deep 

In small wood with 
dense vegetation ; 
300-400 yards from 
nearest house 

15.6.1919. Many 
larvae and pupae of 
O. geniculatus. 

27.9.1919. Few 
larvae of O. geni¬ 
culatus found. 
Adult <£ A. plum¬ 
beus sheltering in 
rot-hole. 

Aigburth Vale, L’pool 
(By kind permission 
of R. B. Miller, 
Esq.) 

Beech 

{Fagus silvatica) 

12 ft. 

16 x 6 x 42 deep 

In grounds of house 
situated near main 
road 

26.7.1919. Numerosj 
larvae of A. plsm- 
beus. 

Storeton, Chcs. 

. 

Beech 

{Fagus silvatica) 

3 ft. 

4x6x12 deep 

In small open and 
wind swept copse 
with little under¬ 
growth. Nearest 
houses, 200-300 
yards distant 

1 

8.6.1919. Many 
larvae and pupae d 
A. plumbeus and 

O. geniculatus 
found. 

— .9.1919. Com¬ 
paratively few and 
mostly small larvae 
of A . plumbeus 
present. 

Bidston, Ches. 

No. i. 

Sycamore 
{Acer pseudo - 
platanus) 

4 i ft- 

7x9x9 deep 

In waste ground on 
outskirts of village 

14.12.1919. Numcrw-i 
very small larvae of 
A. plumbeus. 

No. 2 . 

Do. 

lift. 

10 x 14 x 10 deep 

In small plantation 
bordering road on 
outskirts of village 

14.12.1919. Nunfaca 
very small larvae 

A. plumbeus and i 
f ew large larvae of 
O. geniculatus. 

No. 3. 

Do. 

3 * ft- 

12 x 8i x 12 deep 

do. 

do. 

No. 4 . 

Do. 

3 ft. 

17 x 17 x 20 deep 

In meadow bordering 
road on outskirts 
of village 

24.1.1920. Many 
small larvae of | 

A. plumbeus. 

No. s. 

Do. 

5 ft- 

12 x 17 x 21 deep 

do. 

do. 

No. 6 . 

Do. 

li ft. 

11 x 9 x 6 deep 

do. 

! do. . 
























43i 

TABLE l—continued. 


Details of the breeding-places observed.* 


Locality 

Kind of Tree 

Height of 
aperture 
above ground 

Dimensions of 
rot-hole, in inches 
(approx.) 

Situation 

Remarks 

Westmorland. 

Grasmere 

(By kind permission 
of Sir Francis C. 
Danson) 

Oakf 

( Quercus robur) 

5 * ft. 

24 x 8 x 42 deep 

In private grounds, 
close to house 

6.7.1919. Numerous 
larvae and pupae 
of A. plumbeus. 
Aperture almost 
blocked by a thick 
layer of soil in 
which a large fern 
was growing. 

WALES. 
Flintshire. 
Prestatyn . 

Sycamore 
{Acer pseudo - 
platanus ) 

4i ft. 

12 x 12 x 12 deep 

In lane 200 yards 
from hotel 

20.12.1919. Many 
small larvae of 

A, plumbeus. 

Denbighshire. 

Vbergele. 

No. 1. 

Sycamore 
{Acer pseudo- 
platanus) 

1 

4 ft. 

— 

In school grounds ; 

50 yards from 
houses 

22.12.1919. Small 
to medium size 
larvae of A. plum¬ 
beus. 

No. 2 

Wych Elm 
{Ulmus montana) 

1 

1 

i 5 ft. 


Between two cottages 
and about 10 yards 
from each 

22.12.1919. Small to 
medium size larvae 
of A. plumbeus. 
Occupants of cot¬ 
tages* complain of 

4 bites from buzzing 
gnats in summer, 
which raise swellings 
on arms and legs/ 

langollen . 

No. 1 

Horse-chestnut 
{Aescuius hippo- 
castanum) 

20 ft. 

10 x 6 x 24 deep 

In private grounds, 

50 yards from 
house 

3.1.1920. Small 
larvae of A. plum¬ 
beus numerous. 
Altitude 400 ft. 

No. 2 . 

Do. 

10 ft. 

12 x 12 x 24 deep 

In private grounds, 

10 yards from 
house 

do. 

Carnarvonshire 

nfairfechan 

By kind permission 
of Mervyn T. 
Archdale, Esq., 
M.D.) 

Elm 

{Ulmus campestris) 

lift. 

1 

X 

1 

X 

00 

Q. 

s 

"O 

In small wood with 
fairly dense vegeta¬ 
tion : bordering 
cricket ground 

— .9.1919. Several 
pupae and numer¬ 
ous larvae of 
various sizes of 

A. plumbeus. 

rRE LAND. 

Co. Dublin. 

gstown . 1 

'0. 1 

1 

Sycamore 
{Acer pseudo - 
platanus) 

lift. 

[ 

12 x 12 x 12 deep 

In private grounds ; 
several houses 
within a radius of 

50 yards 

30.12.1919. Small 
and medium size 
larvae of A. plum¬ 
beus numerous. 

0. 2 . 1 

Do. 

13 ft. 

16 x 9 X 18 deep 

In garden ; house 
within 15 yards 

do. 

rstown District ... 
y kind permission 
of M. V. Blacker 
Douglass, Esq.) 

Do. 

8 ft. 

10 x 10 X4 deep 

In private grounds ; 
500 yards from 
house 

27.12.1919. Few 
small larvae of 

A. plumbeus. 

--- 

icy .; 

bserver Mrs. 

> top ford Douglas) 

Do. 


; 

Nearest house 300- 
400 yards distant 

26.1.1920. Small 
larvae of A. plum¬ 
beus. 


* Scotland. Breeding-places see footnote, p. 446. 
-f- Al*o in an Ash tree (Fraxinus excelsior) 14.2.1920. 




432 


sometimes necessary when the aperture is small. Many rot-holes of 
course are conspicuous and can be detected several yards away, but 
the apertures of others are small or at considerable heights above 
the ground or are so well concealed by rotten pieces of branch or 
ferns that most careful examination is necessary. The location of 
breeding-places is distinctly easier in winter. We have been obliged 
to confine ourselves to the examination of holes situated not more 
than twenty feet from the ground, but even with such a limit it is 
extremely difficult to decide that breeding-places, potential and 
actual, do not exist in any given locality; and we believe that the 
proof of their non-existence within a given area will necessitate the 
closest scrutiny of every tree within that area. 

Larval Habits 

The larvae of A. plumbeus were, in all probability, first taken by 
Grassi (1901) in Italy, but it is abundantly clear that this author 
associated them with A. bifurcatus and regarded the peculiar type 
_ of breeding-place as occasional for this latter species. Eysell (1912) 
first described the larvae, his specimens being reared from eggs laid 
by captive females. Meinert’s (1886) description of the larva of 
A. nigripes clearly applies to that of A. bifur catus as may readily 
be perceived from his figure. To Galli-Valerio and de Jongh 
(1912 2 and 1913) must be given the credit of finding them in nature 
and of recognising the great selection shown by the species in its 
choice of a breeding-place. 

Their movements and habit of living mainly at the surface of 
the water appear, judging by laboratory specimens, to be typically 
Anopheline. Galli-Valerio and de Jongh (1913), however, state 
that they require very little air, seldom show themselves at the 
surface and remain chiefly at the bottom among the debris. We 
have not observed this to be the case and think it probable that these 
authors were misled by the rapidity with which the larvae descend 
when disturbed. On the slightest provocation the larvae move down¬ 
wards, the small ones wriggling actively, the large ones usually after 
a few strong jerks of their bodies sinking motionless to the bottom 
where they lie in any position among the debris. Larvae, particu¬ 
larly older ones, will remain apparently dead at the bottom for 
considerable periods; we have watched them lie thus for twenty 
minutes or half-an-hour. 



433 


The main food of the larvae is as yet undecided. Christophers 
(1916) and Edwards (1916) maintain that they are carnivorous or 
semi-camivorous, existing largely upon insects which fall into the 
water. The larvae of A . barberi and A. culicifortnis moreover show 
strong cannibalistic tendencies and it is probable, therefore, that 
the above opinion is correct. We have observed that both 
A. plumbeus and O. geniculatus larvae are greatly attracted by the 
dead and floating bodies of ‘rat-tailed maggots’ of which they 
ultimately leave only the tips of the syphon-tubes. Eysell (1912) 
successfully reared adults from the egg by supplying the larvae with 
bacterial scum from hay infusion. 

The stage in which hibernation takes place has been variously 
surmised, and although Galli-Valerio (1917 1 ) found larvae in tree- 
holes the water of which was covered with ice, no decision on the 
matter seems to have been reached. Eckstein (1918*) discussing this 
question states that it is unlikely that they over-winter as larvae, as 
during the winter the tree-holes either dry out or the water is frozen 
into a solid block; for this reason, also, he has never found hiber¬ 
nating larvae in spite of having repeatedly searched known breeding- 
places. We fail to follow this author’s statements and are convinced 
that A. plumbeus does hibernate in the larval stage. We have 
obtained young larvae on numerous occasions in December, 
January and February, and have found no evidence of the holes 
drying out during these months. That the larvae can revive after 
the water in which they live has been frozen solid for short periods 
is shown by the following experiments (Table II). 

We can offer no explanation of the results obtained in Experi¬ 
ment 5. In all the experiments the larvae were contained in water 
taken with them from the rot-holes. In Experiments 3-9 all the 
larvae after subjection to the low temperature were encased in blocks 
of ice, and even when the surface layers of water remained liquid, 
they were embedded in the ice below. This appeared to be due to 
the paralysing effect of the low temperature (acting from below) 
causing them to sink to the colder layers of the liquid which shortly 
after became frozen. In most cases the larvae at the end of the 
experiments resumed their activities remarkably quickly, and many, 
particularly the small ones, showed signs of movement directly their 
bodies, or portions of their bodies, were freed from the ice. 



434 


TABLE II. 
Freezing Experiments 


Experi¬ 

ment 

i 

Species 

Description of 
larvae used 

Amount 1 
of water ! 
in larval | 
container ' 

1 

1 

Tempera¬ 

ture 

of water 
before 
freezing 

Time 

immersed in 
freezing 
mixture 
(Temp, of 
mixture 
-4 0 C. to 
-2°C) 

Result of 
immersion 
on water 
containing 
larvae 

Condition of 
larvae 20 hours 
after removal of 
container from 
freezing mixture 

Remade 

i 

! 

i 

A. plumbeus 

Five small and 
one apparently 
half-grown 

i 

1 

5 c c - ; 

i 

if c. 

5 minutes 

i , 

No ice 
present 

All active and 
apparently 
unaffected 


2 

Do. 

do. 

do. 

do. 

10 minutes 

do. 

do. 


3 

Do. 

Fout small 

do. 

do. 

15 minutes 

Lower 

layers 

containing 

larvae 

frozen 

solid 

do. 

One larva 
remained 
motionless 
for at least 

1 one hour after 
removal of 
container fron 
mixture 

1 

4 

Do. 

Five small and 1 
one apparently 
half-grown 

* do. 

do. 

20 minutes 

Frozen 

solid 

do. 


5 

Do. 

do. 

• 

do. 

do. 

25 minutes 

do. 1 

i 

i 

One small larva 
active ; rest 
apparently 
dead, but j 

showing slight 
movement 
under bino¬ 
cular 

! The small lam 

1 regained 

activity im¬ 
mediately cm 
thawing. 
dead 40 bstr* 
after reimmi 
of contain* 
♦rom mixrn-t 

6 

Do. 

do. 

C.C. 

do. 

3c minute? 

do. 

All active and 
apparently 
unaffected 

! 

7 

Do. 

■ 

10 C.C. 

4- c c.c. 
debris 

do. 

do. 

do. 

do. 


8 

0 . oemculatus 

Four half-grown 
and one 
apparently 
mature 

15 c.c. 

do. 

do. 

1 do. 

1 

1 

do.. 

‘ 

i 

i 


9 

Do. 

One half-grown 
and one nearly 
mature 

15 C.C. 

do. 

1 

do. do. do. 

1 

1 ! 

1 





435 


In connection with the above experiments Howard, Dyar and 
Knab’s (1912) remarks concerning the larvae of the pitcher-plant 
mosquito, Wyeomyia smithii , are of interest; they state that ‘ the 
larvae survive complete freezing up of the water in the leaves of the 
plant and complete their development in the following season.’ 

The duration of the larval stage in summer appears to be about 
four weeks. Eysell (1912) gives the length of this stage as 28 days 
and the cycle from egg to adult as 33-35 days, and Christophers 
(1916) obtained the full evolution in about four weeks. The larvae 
observed by both these authors received an ample supply of food. 
Galli-Valerio and de Jongh (1913) give the minimum time for the 
completion of the cycle as 30 days and state that the larval period 
is often prolonged, so that the metamorphosis from egg to adult 
may occupy as long as eleven months. The factor of hibernation 
entered into the last named period, but in cases where it did not act 
and the larval stage was prolonged the cause may have been a 
deficient or unsuitable food supply. These observers make no partic¬ 
ular reference to the food provided, and the inference therefore is 
that the larvae were merely placed in jars with water and debris from 
the rot-hole, and such conditions, according to Edwards (1916) do 
not favour rapid development. 

The larvae of A. plutnbeus are often found associated with those 
of the Aedine mosquito, Ochlerotatus geniculatus , Ol., with which 
they appear to live in amity. Recently Macgregor (1919 1 & 2 ) has 
found them in Epping Forest with the larvae of a previously 
undescribed species of Orthopodomyia and apparently also with 
those of Siegomyia fasciata , Fabr.* Other dipterous larvae not 
•infrequently occurring in the rot-holes are those of various 
Chironomids and the ‘ rat-tailed maggots * of Myiatropa florea , L. t, 
and Helophilus pendulus , L. 

Distribution of A. plumbeus in the British Isles 

The known distribution of this mosquito in this country is shown 
in Map 2. At present records from about fifty localities only are 
available, and most of these are isolated and due to the capture of 

* The larvae of this mosquito were not actually observed by Macgregor, but two males 
emerged from the tank in which the tree-hole species were kept. 

t Identification kindly confirmed by Major E. E. Austen of the British Museum. 


436 


occasional blood-seeking females. Such records spread over a long 
period of time, and give no clue to the abundance of the species. 
Much attention has been given to our other indigenous Anophelines— 
A. maculipennis and A. bifurcatus —and valuable information has 
been obtained regarding their bionomics and distribution, but 
A. plumbeus > on account of its apparent rarity, has been greatly 
neglected. Even in places where it was known to occur, no serious 
attempts to discover its habits appear to have been made, and it 
was taken for granted that in these it resembled the other two 
species. Edwards (1916), following Galli-Valerio and de Jongh’s 
(1912 and 1913) work in Switzerland, discovered the breeding- 
places of A . plumbeus at Burnham Beeches in Buckinghamshire, but 
until last year not more than three or four additional records of this 
nature had been made. Our investigations began in June, 1919, 
and the direct results, so far as this line of research is concerned, 
are shown in the Maps 1 and 2 and in Table I. We have found 
breeding-places in every locality which we have searched, and 
although no attempts at intensive surveys have been made, and no 
idea of the numerical distribution of the mosquito in any locality 
obtained, we are convinced that the species is widely and commonly 
distributed throughout the country. 

Occurrence of A. plumbeus in the Liverpool District 

With reference to the occurrence of the adults of this species in 
the Liverpool district, Newstead informed Scott (quoted by Lang 
(1918)) that A. plumbeus was relatively scarce. Beyond this, the 
only records from Lancashire are those of Theobald (1910, and in 
Lang (1918)) from Manchester. Map 1 shows that we have 
discovered a number of breeding-places of this species at varying 
distances from the centre of the city and at approximately equal 
distances from each other. But it must not be supposed that no 
breeding-places exist in the intervals. In all probability they do, but 
time did not allow of a more extensive survey, and we were obliged to 
limit ourselves to particular localities. At the commencement of 
the investigations we naturally selected spots which we thought, 
from the nature of the country, would be most likely to prove 
satisfactory. But as success attended our search and the diffuse 
nature of the distribution of this mosquito was realised, we only 



437 



Map I 


LIVERPOOL DISTRICT 

0 indicates breeding places 
of Anofbeks flumbeus , Steph. 






438 


chose such places as would, by their direction, enable us to 
complete as far as possible the cordon of breeding-places round the 
town. In one of these districts the outlook was most unpromising, 
as trees were by no means numerous; yet the result was successful, 
and a breeding-place (several likely rot-holes too high for us to 
examine were also seen) was found within two hours. At first we 
examined places situated from five to eight miles away, but 
subsequently we reduced the distance. Within the four-mile limit 
the work was more difficult, as most of the trees are enclosed in 
private property; nevertheless on several occasions breeding-places 
were found. 

From the above remarks it will be seen that A. plumbeus is not 
uncommon in the Liverpool area, that it occurs widely distributed 
round the city and within the four-mile limit. 

Possible connection of A . plumbeus with the Spread of 
Malaria 

The importance of A. plumbeus as a natural carrier of malaria 
cannot yet be estimated. Far more detailed information regarding 
its bionomics and distribution is required before any definite 
statement can be made. But in our opinion the matter is not to be 
dismissed so lightly as is being done; on the contrary, A. plumbeus 
is worthy of serious consideration in this respect. 

In country districts or woodlands much frequented by the public 
the possibility of the dissemination of the disease by this mosquito can 
be readily appreciated; for when old trees or trees whose preserva¬ 
tion has been neglected are numerous the opportunities for breeding 
will be greater, and the output of adult mosquitoes increased. But 
in towns or villages not situated in the immediate neighbourhood 
of large woods the question is more complex. It will be urged that 
A . plumbeus is a‘wild’ species, and that in such places it is 
extremely rare and consequently of little or no importance. This 
may be so, but neither the degree of its domestication nor its 
numerical distribution can be measured solely by the frequency with 
which adults are observed in man’s dwellings. The extent to which 
it occurs in and around the larger towns can only be decided by 
careful intensive surveys for breeding-places. The adult habit of 
resting in the rot-holes renders any conclusions drawn from an adult 



439 


census elsewhere entirely misleading. From 1910 to 1916 a few 
isolated specimens only of A . plumbeus had been found in Northern 
India, but Christophers (1916), after his discovery of the breeding- 
places at Simla and his subsequent observations, stated that 
1 there can be no doubt that in the particular area investigated 
Anopheles (in the form of A. barianensis*) must have been 
quite numerous/ It seems to us not improbable that the highly 
selective and unusual habits of a mosquito such as A. plumbeus may, 
at least in some cases, have a direct bearing upon the condition 
known as ‘ Malaria sine Paludism/ where malaria may occur in a 
locality which appears relatively free from Anophelines. It has 
been repeatedly observed that marshiness and the abundance of 
Anophelines are not always in proportion to the extent and severity 
of a malaria epidemic; Celli (1903), in fact, states that in Basilicata 
in Italy, and in Sicily, this proportion is inverse, the area of marsh 
being very small and sometimes limited to the beds of torrents or 
streams and Anophelines scanty, while malaria is common and of 
a severe type. 

It has been shown (cf. p. 429) that the breeding-places of 
A. plumbeus may occur in more or less isolated trees, and when 
such trees are situated within a few yards of a house (cf. Table I, 
Fazakerley, Grasmere, Abergele No. 2, Llangollen No. 2, and 
Kingstown Nos. 1 and 2) it is practically certain that the females 
in their search for blood will frequent this house with considerable 
regularity. 

These considerations make it desirable to investigate fully 
the immediate surroundings of any houses to which the occurrence 
of cases of malaria seems peculiarly restricted, with a view to 
discovering the breeding-places of A. plumbeus. 

ACKNO WLBDGM ENTS 

We are indebted to Professor W. Ramsden for analysing two 
samples of rot-hole water submitted to him. Our thanks are also 
due to Mrs. Stopford Douglas for obtaining and sending larvae 
from Killiney, Co. Dublin, and to those gentlemen, named in 
the table of breeding-places, who have courteously granted us 
permission to examine the trees in their private grounds. 

* An barianensis is synonymous with A . plumbeus (sec p. 422). 



PRELIMINARY REPORT BY PROFESSOR W. RAMSDEN 
ON LIQUIDS FROM CAVITIES OF HOLLOW TREES 

Specimen i. Filtrate of mixed samples of liquids in all 
of which A. PLUMBEUS was breeding. 

Inodorous. No froth when shaken. Brown (porter) colour with 
slight greenish fluorescence; spectroscopically, general absorption of 
the blue end, but no definite band. Specific gravity: 1004. 

Total solids from 100 c.c. evaporated to dryness on water-bath 
0 960 grams. 

Total Nitrogen by Kjeldahl method: 00053 grams in 100 c.c. 

Reaction : markedly alkaline to litmus. 

Gives off CO a with mild effervescence when acidified, therefore 
contains Carbonates. 

Dry residue chars strongly when heated sufficiently; smell does 
not suggest either proteins or carbohydrates. 

Ash obtained rich in Calcium, Potassium and Phosphates, and 
contains also Sodium, Iron, Magnesium, Chlorides and Sulphates. 

Surface-tension: markedly less than water. 

The enzyme ‘ catalase ’ is absent. 

A doubtful Molisch’s reaction is obtainable, although a small 
quantity of a Furfural is given off when the liquid is boiled with 
60 per cent. H a S 0 4 (anilin acetate turned pink): therefore some 
carbohydrate grouping in small quantity is presumably present. 

No fermentation with yeast. No colour with Iodine solution. 
No reduction of Fehling’s or Nylander’s solutions. 

The brown pigmented organic matter is precipitated on adding 
any strong mineral acid or oxalic acid, but not by acetic acid. It 
is also salted out by saturation with ammonium sulphate. It gives 
a strong ‘ xanthoproteic ’ reaction. It is not a protein. 

SPECIMEN 2.* Filtrate of sample in which no larvae were 

found. 

Specific gravity was greater: 1009, but in every qualitative 
respect the statements made above for the first liquid are equally 
true for the second. 


* See footnote on p. 428. 




LITERATURE 


Alcocic, A- (1911). Entomology for Medical Officers, London, p. 70 

Austen, E. E. (1906). Illustrations of British Blood-sucking Flies. London, p. 18. 

Bacot, A. (1918). Mosquitoes and the danger of Malaria in England. Essex Naturalist. Vol. 18, 
pp. 241-263. 

Bastianelli, G., Bignami, A., and Grassi, B. (1898). Zoologia mcdica—Coltivazione delle 
semilune malariche dell* uomo nelT Anopheles claviger , Fabr. (sinonimo : Anopheles 
maculipennis , Meig.) Atti della Reale Accad. dei Lincei . VoL 7, pp. 313-314* 

Bczzi, M. (1892). Contribuzione alia fauna Ditterologica della provincia di Pavia. Bull. Soc. 
Ent. Ital. Anno. 24, p. 133. 

Black lock, B., and Carter, H. F. (1919). The Experimental Infection of Anopheles plumheus , 
Hal., with Plasmodium vivax. (Sporozoites in Salivary Glands). Preliminary Note. 
Ann. Trop. Med. & Parasitol. Vol. 13, pp. 187-188. 

Blanchard, C. (1918). Sur un nouveau type larvaire du groupe des Anophelines. Bull. Soc. 
Path. Exot. T. 10, pp. 669-677. 

Blanchard, R. (1905). Les Moustiques. Histoire naturelle et medicale Paris, pp. 164 and 166. 

- (1918). Bull, de VAcadimie de Medicine. August. (Note regarding the identity of 

the larvae obtained by C. Blanchard, vide supra). 3e S6r. T. 80, p. 146. 

Celli, A. (1903), La malaria in Italia durante il 1902. Atti della Soc ., per gli Studi della 
malaria . Vol. 4, p. 551. 

Christophers, S. R. (1916). An Indian tree-hole breeding Anopheles , A. harianensis , James= 
A. ( Coelodiazesis ) plumheus , Haliday. Ind. Journ. Med. Res. Vol. 3, pp. 489-496. 

Christophers, S. R., and Khazan Chand (1916). A tree-hole breeding Anopheles from 
Southern India : A. ( Coelodiazesis) culiciformis , Cogill. Ind. Journ. Med. Res. Vol. 3, 
pp. 638-645. 

Coquillett, D. W. (1903). A new Anopheles with unspotted wings, Canad. Ent , Vol. 
35 > P* 3 I0 « 

Cordier, E. (1918). Capture en Argonne d ’Anopheles nigripes , Staeger 1839, espece nouvellc 
pour la faune fran^aise. Bull. Soc. Path. Exot. Tome 11, pp. 726-727. 

Donovan, C. (1901). Anopheles in Ireland. B. M. J. Vol. 2, p. 14. 

Dyar, H. G. (1918). Notes on American Anopheles. Insecutor Inscitiae Menstruies. Vol. 6, 
pp. 141-151. 

Dyar, H. G., and Knab, F. (1906). The larvae of Culicidae classified as independent 
organisms. Journ. N. T . Ent. Soc % Vol, 14, p. 177. 

Eckstein, F. (1918 1 ). Die Ueberwinterung unserer Stechmucken. Biol. Central. Bd. 38, 

pp- 530-536 

- (1918*). Zur systematik der einheimischen Stechmucken. 1. Vorlaufige Mitteilung : 

Die Weibchen. Cent./. Bakt. Parasit. u. Infektionskr. ite Abt. Orig. Bd. 82, pp. 
57-68. 

-(1919). Zur systematik der einheimischen Stechmucken. 2. Vor. Mitt: Die Larven. 

Ibid. Bd. 83, pp. 281-294. 

Edwards, F. W. (1912). Notes on the British Mosquitos (Cuiicinae). The Entomologist. Vol. 
* 45 i PP* , 93 " I 94 * 

-(1916). Notes on Culicidae, with descriptions of new species. Bull. Ent. Res. Vol. 

7, p. 201. 

Eysell, A (1912)* Cyclopborus {Anopheles) nigripes , Staeger (nov. gen.) Arch. f. Scbijfs u. 
Tropen Hyg. Bd. 16, pp. 421-431. 

- ( I 9 I 3 )* Die Stechmucken. Handbucb der Tropenkrankbeiten. Leipzig. Bd. 2, pp. 

44 * 94 - 

Ficalbi, E. (1896). Revisione sistematica della famiglia delle Culicidae Europee. Firenze, 
pp. 99, 109 and 217. 

-(1899). Venti specie di Zanzare (Culicidae) italiane classate, descritte e indicate secondo 

la loro distribuzione corologica. Bull. Soc. Ent. Ital. Anno. 31, pp. 46-234. 



44* 


Galli-Valerio, (19151). Studi e ricerche sui Culicidi. 11 a Mem. La Malariologia. Vol. 8, 

pp. 141-143* 

-(1915*). Beobachtungen uber Culiciden. Cent. f. Baku Parasiu u. Infektionskr. 

ite. Abt. Orig. Bd. 76, pp. 260-261. 

- (*9*6). Beobachtungen fiber Culiciden. Cent.f. Baku Parasiu u. Infektionskr. ite. 

Abt. Orig. Bd. 78, pp. 90-96. 

- (* 9 * 7 1 ). Beobachtungen fiber Culiciden. Ibid. Bd. 79, pp. 139-143. 

- (1917*). La distribution geographique des Anoph 61 ines en Suisse au point de vue du 

danger du formation de foyers de Malaria. Bull. Office Internat. d*Hyg. Publique , Paris. 
T. 9, pp. 1566-1582. 

Galli-Valerio, B., and Rochaz de Jongh, J. (1903). Studi e ricerche sui Culicidi dei generi 
Culex e anopheles. Atti della Soc. per gli Studi della malaria. Vol. 4, p. 44. 

- (1912 1 ). Beobachtungen fiber Culiciden und Mitteilung fiber das Vorkommen von 

Pblebotomus papatasi , Scop, im Kan ton Waadt (Schweiz). Cent. f. Baku Parasiu u. 
Infektionskr. ite. Abt. Orig. Bd. 63, pp. 222-227. 

-(1912 9 ). Studi e ricerche sui Culicidi. 8a Mem. Atti della Soc. per gli Studi della 

malaria. Vol. 13, pp. 1-5. 

-(*913). Beobachtungen fiber Culiciden. Cent. /. Baku Parasiu u. Infektionskr. 

ite. Abt. Orig. Bd. 67, pp. 472-478. 

-(19141). Beobachtungen fiber Culiciden. Ibid. Bd. 72, pp. 529-53*. 

-(1914*). Studi e ricerche sui Culicidi. 9a Mem. Atti della Soc. per gli Studi della 

Malaria. Vol. 14, pp. 1-5. 

-(1915). Studi e ricerche sui Culicidi. ioa Mem. La Malariologia. Vol. 8, 

pp. 9-11. 

Giles, G. M. (1900). A Handbook of Gnats or Mosquitoes. London. 1st ed., p. 175. 
-(1902). Ibid. 2nd ed., p. 330. 

- (1904). A revision of the Anophelinae being a first supplement to the second edition 

of ‘ A Handbook of Gnats or Mosquitoes.' London, p. 23. 

Grass 1, B. (1898 1 ). Zoologies medica—Rapporti tra la malaria e peculiari insetti (zanzaroni 
e zanzare palustri). Atti della Reale Accad. dei Lincei. Vol. 7, pp. 163-172. 

- (1898 9 ). Zoologica medica—La malaria propagata per mezzo di peculiari insetti. 

Ibid, pp. 234-240. 

- (1898 3 ). Zoologica medica—Rapporti tra la malaria e gli artropodi. Ibid , pp. 314-315. 

- (1899). Ancora sulla malaria. Ibid. Vol. 8, p. 559. 

- (1901) Die Malaria. Studien eines Zoologen. ite vermebrte Auflage. Jena, pp. various. 

- (1903). Documenti riguardanti la storia della scoperta del modo di trasmissione 

della malaria umana. Milano, p. 96. 

Grass 1, BL, Bignami, A., and Bastianelli, G. (1899*). Ciclo evolutivo delle semilune nell’. 
Anopheles claviger ed. altri studi sulla malaria dalT Ottobre 1898 al maggio 1899. Atti 
della Soc. per gli Studi della Malaria. Vol. I, pp. 14-27. 

-(1899*). Resoconto degli studi fatti sulla malaria durante il mese di gennaio. 

Rendiconti della Reale Accad. dei Lincei . Vol. 8, Feb. 5. 

- (*899*). Ulteriori ricerche sulla malaria. Ibid. Vol. 8., May 7. 

Grunberg, K. (1910). Diptera Zweiflfigler. Brauer: Die Sfisswasserfauna Deutschland*. 
Heft. 2 A., p. 83. 

Haliday, A. H. (1828). Notice of Insects taken in the North of Ireland. Zool.Journ. Vol. 3 

- (*833). Catalogue of Diptera occurring about Holy wood in Downshire. EnU Mag. 

Vol. 1, pp. 148 and 151. 

Hermes, W. B. (*915). Medical and Veterinary Entomology. New York, p. 99. 

Hesse, E. (1918). Sur la presence dans le Dauphin^ de 1 ' Anopheles nigripes , Staeger. Arch. 
Zool. Exph. Nous et Revue. Paris, T. 57, pp. 32-35. 

Hindle, E. (1914). Flies in Relation to Disease. Blood-sucking Flies. Cambridge, pp. 81 
and 102. 



443 


Howard, L. O., Dyar, H. G., and Knab, F. (1912-1917). The Mosquitos of North and 
Central America and the West Indies. VoL 1, p. 417. Vol. 4, pp. 1035-1038. 

James, S. P. (1911). Anopheles barianensis , James. A monograph of the Anopheline Mosquitoes 
of India. 2nd edition. Calcutta, pp. 55 and 76. 

- ( I 9 I 7). Note recording the proof that Anopheles tnaculipennis is an efficient host of 

the Benign Tertian parasite in England, Journ. RrA.M.C . Vol. 29, p. 615. 

Jarvis, F. E. (1919). On the occurrence of the immature stages of Anopheles in London. 
Ann. Applied Biology. Vol. 6, pp. 40-47. 

Lang, W. D. (1918). A map showing the known distribution in England and Wales of the 
Anopheline Mosquitoes with explanatory text and notes. British Museum {Nat. Hist.) 
London, pp. 48-52 and 63. 

Loew, H. (1845). Dipterologische Beitrage. Posen, p. 4. 

Langeron, fa. (1918). Presence de VAnopheles nigripes , Staeger, 1839 dans la region parisienne. 
Bull. Soc. Path. Exot. T. 11, p. 728. 

MacGregor, M. E. (1919 1 ). On the occurrence of Stegomyia jasciata in a hole in a beech tree 
in Epping Forest. Bull. Ent. Res. Vol. 10, p. 91. 

- ( I 9 I 9 m )- A new mosquito of the genus Ortbopodomyia from a beech tree-hole in England. 

Journ. R.A.M.C. Vol. 33, pp. 451-454. 

Martini, E. (1915). Uber drei weniger bekannte deutsche Kuliziden: Aides ornatus, Mg. 
Mansonia ricbiardii, Fic. und Anopheles ( Coelodiazesis ) nigripes , Staeger. Arch. f. Scbijfs . 
u. Tropen. Hyg. Bd. 19, pp. 585-607. 

Meinert, F. (1886). De eucephale Myggelarver. Sur les larves euctphales des Dip teres. 
Leurs moeurs et metamorphoses. Danske videnskabeme Selsk. Skrift. Bd. 3, pp. 
373 - 493 - 

Mitzmain, M. B. (1916). Anopheles infectivity experiments. U. S. P. H. S. Pub. Health Rep. 
Washington. Vol. 31, p. 2325. 

Nuttall, G. H. F. (1899-1900). Neuere Forschungen uber die Rolle de Mosquitos bei der 
Verbreitung der Malaria. Cent. f. Baku, Parasit. u. Injektionskr. ite Abt. Bd. 25, 
pp. 877-881. Bd. 26, pp. 140-147 and Bd. 27, pp. 218-225. 

Nuttall, G. H. F., Cobbett, L., and Strangeways-Pigg, T. (1901). Studies in Relation to 
Malaria, I. The geographical distribution of Anopheles in Relation to the former 
distribution of ague in England. Journ. Hyg. Vol. 1, pp. 4-44. 

Nuttall, G. H. F., and Shipley, E. (1901). Studies in Relation to Malaria II. The Structure 
and Biology of Anopheles {Anopheles maculipennis). The Egg and Larva. Ibid. Vol. 1, 
. PP- 45 - 73 - 

Osten-Sacken, C. R. (1878). Cat. Dipt. N. America. 2nd ed., p. 19. 

Parsons, A. C., and Brook, G. R. (1919)- The mosquito problem in Britain : Suggestions 
for a winter campaign against the important mosquitoes, with notes on Insecticides. 
Journ. R.A.M.C. Vol. 32, pp. 1-23. 

Patton, W. S., and Cragg, F. W. (1913). A Text-book of Medical Entomology. Calcutta, 
p. 241, 

Peju, G., and Cordier, E. (1918). Epid6mie palustre et conditions de l'Anophtlisme en 
Argonne. C. R. Soc. Biol., Paris. T. 81, pp. 1039-1041. 

- (*919- Paludisme et Topographic Anoph&ienne en Argonne a propos d’une 

6pidemie de paludisme autochtone. Bull. Soc. Path. Exot., Paris. T. 12, pp. 23-24. 

Schiner, J. R. (1864). Fauna Austriaca. Die Fliegen (Diptera). Wien. Bd. 2, p. 625. 

Schneider, P. (1914)* Beitrag zur Kenntnis der Culiciden in der Umgebung de Bonn. Ver- 
hand. Naturhist. Ver Preuss. Rheinlande u. Westfalens, Bonn. Bd. 70, pp. 1-54. 

Schoo, H. J. M. (1902). La Malaria in Olanda. Atti della Soc. per gli studi della Malaria. 
Vol. 3, pp. 189-208. 

Smith, J. B. (1906). Report of the mosquito investigation in 1905. Rep. Ent. Dept., New 
Jersey Agric. Expt. Sta.,for the year 1905. pp. 671-675, 

Staeger, C. (1839). Systematik Fortegnelse over de hidtil i Danmark fundne Diptera. Krojer: 
Naturhistorisk Tidskrift. Bd. 2, p. 552. 



444 


Stephens, J. F. (1828). Note on the foregoing paper (Haliday, ut supra ) with a description 
of a new species of Anopheles {Anopheles grisescens). Zool. Journ. Vol. 3. 

- (1829). A systematic catalogue of British Insects. London. 

Stephens, J. W. W., and Christophers, S. R. (1908). The practical study of Malaria. London. 
3rd. ed., p. 158. 

Strobl, G. (1894). Die Dipteren ^on Steiermark. Mittbeil. Natur. ver . J. Steiermark . Bd. 31, 
p. 200. 

Theobald, F. V. (1901). A Monograph of the Culiddae of the World. Vol. 1, p. 202. 

- (1905). Culicidae. Wytsman : Genera Insectorum. Bruxelles 26me Fasc.. p. 7. 

- (1908). Mosquitoes and Peat. Nature , Oct. 15, pp. 607-608. 

-(1910). A Monograph of the Culicidae of the World. Vol. 5, pp. 12 and 13. 

Walker, F. (1856). Insccta Brittannica. Diptera. VoL 3, p. 249. 

Wingate, W. J. (1906). A preliminary list of Durham Diptera, with analytical tables. Trans. 
Nat. Hist. Sec. of Northumberland , Durham and Newcastle-upon-Tyne. Vol. 2, p. 62. 

Wright, M. J. (1901). The resistance of the larval Mosquito to cold. Notes on the habits 
and life-history of mosquitoes in Aberdeenshire. B. M. J. VoL I, pp. 882-883. 

Wulp, F. M. VAN DER (1877). Diptera neerlandica. De Tweevleugeligc Insecten van Neder 
land. *s Gravenhague. Deel. i, p. 331. 

Zettkrstedt, J. G. (1850). Diptera Scandinaviae disposita et descripta (Lundac). Insecta 
lapponica. Lipsia. Tomus 9, p. 3467. 


(1918). Reports to the Local Government Board on Public Health and Medical Subjects. 
(New series, No. 119). 

Reports and papers on Malaria contracted in England in 1917. London. 

(1919). Ibid. (New series, No. 123). Reports and papers on Malaria contracted in England 
in 1918. London. 


Note. —(cf. p. 422)—A paper by B. Prashad on ‘ The description and life-history of a 
new species of Anopheles that breeds in holes in trees* {Rec. Ind. Mus n VoL 15, iii, 1918, 
pp. 123-127) has come to our notice too late for reference in the text. This author describes 
Anopheles annandalei , from a male, bred from a larva collected from a tree-hole in the 
Eastern Himalayas. The adult possesses spotted wings and a conspicuous tuft of scales on 
each of the hind femora and thus superficially resembles A. asiatiea , Leic. Prashad indeed 
groups it with this species and with A. bar biros tris, Wulp, and A. wellingtontanus , Alcock, 
but the excellent description and figure of the larva of A. annandalei given by this writer 
demonstrate conclusively that it is a member of the sub-genus Coelodiazesis. 




446 

REFERENCES TO LOCALITY RECORDS SHOWN IN 

MAP II * 


England. 

1. Penzance and Helford, R., 

Cornwall. 

2. Ugbrooke, Devonshire. 

3. Sidmouth, Devonshire. 

4. Lyndhurst, Hampshire. 

5. New Forest, Hampshire. 

6. Wye, Kent. 

7. fBoxhill, Surrey (F. W. Edwards, 

Esq;, private communication) 

8. Merton, Surrey. 

9. Acton, Middlesex. 

10. Bushey Heath, Hertfordshire. 

11. fCassiobury Park, Watford, 

Hertfordshire 

12. fEpping Forest, Essex 

13. Rochford and Gt. Wakering, 

Essex. 

14. tBurnham Beeches, Buckingham¬ 

shire. 

15. fHitchen, Hertfordshire (F. W. 

Edwards, Esq., private 
communication). 

16. Radwell, Hertfordshire. 

17. ’(■Cambridge (F. W. Edwards, 

Esq., private communication). 

18. Newmarket, Suffolk. 

19. Butley, Suffolk. 

20. Oxford. 

21. Lamport, Northamptonshire. 

22. ■fLongner Hall, S.E. of Shrewsbury, 

.(R. F. L. Burton, Esq., 
private communication). 

23. Attingham Park, Shropshire. 

24. Shrewsbury. 

25. Wood Eaton Manor, 

Staffordshire. 

26. Bakewell, Derbyshire. 

27. Poynton, Cheshire. 


28. Manchester. 

29. fRoby and Knowsley Moss, 

Lancashire. 

30. fAigburth Vale, Liverpool. 
jfHightown, Lancashire 

* 1 * t fFazakerley, Liverpool 

32. ■fStoreton, Cheshire 

33. •j'Bidston, Cheshire 

34. fGrasmere, Westmorland 

Wales.* 

35. ■(•Prestatyn, Flintshire 

36. f Abergele, Denbighshire 

37. Colwyn Bay, Denbighshire. 

38. fLlanfairfechan, Carnarvonshire. 

39. Bettws-y-Coed, Carnarvonshire. 

40. Beddgelert, Carnarvonshire. 

41. * Llangollen, Denbighshire 

42. Llangammarch Wells, 

Brecknockshire. 


Scotland.! 

43. Edinburgh. 

44. Culross, Fifeshire. 

45. Aberdown Woods, Fifeshire 

46. Fife. 

47. Blairgowrie, Perthshire. 

48. Torphins, Aberdeenshire. 

Ireland. 

49. Holywood, Co. Down. 

50. Harold’s Cross, Co. Dublin. 

51. f Kingstown, Co. Dublin 

52. f Killiney, Co. Dublin 

(Mrs. Stopford Douglas). 

53. Courtmacsherry, Co. Cork. 


•This map is based upon that recently published by Lang, and most of the records of 
adult captures have been obtained from his work. 

t Indicates breeding place found. 

X Additional localities:—Trawsmawr, 4 miles north of Carmarthen, September, 1907. 
Prof. J. W. W. Stephens. Three females captured in garden when attempting to bite. 
Scotland: Skibo Castle, Sutherland, 29.2.1920. Breeding place in sycamore tree. Feam, 
Ross and Cromarty, 1.3.1920. Breeding places in beech and scyamore trees, nearest houses 
300-500 yards distant. 




Annals Trop. Med. & Parasttol ., Voi, XI11 


MAP II 



C. Tinting « 5 ^ Co., Ltd., Imp 










EXPLANATION OF PLATES 

Plate X 

Breeding-places of Anopheles plumbeus, Steph., in the Liverpool 
district. 

Fig. i. Rot-hole in sycamore tree (vide record Bidston No. 4 in 
Table I). 

Fig. 2. Rot-hole in sycamore tree (vide Bidston No. 3). 

Fig. 3. Rot-hole in sycamore tree (vide Bidston No. 5). 

Fig. 4. Rot-hole in sycamore tree (vide Bidston No. 2). 



4ntuih Trop. Med, & ParasitolVol, XI11 


PLATE X 



Fic. 4 


C. Tinting <V CoLtd., Imp 


Photographs by Mis* 1 /. Pro:/ 


FlC. I 


* 

Sffw 1 

w •*» 


^i<v >>V#v 

% ■ ti SrV 

'’jraKa* 

jjgX? if? 

rJ 9 fc 








45 © 


Plate XI 

Breeding-places of Anopheles plumbeus, Steph. 

Fig. i. Rot-hole in tree (vide record Kingstown No. 2 in Table I). 

(Photograph by Commander Stopford Douglas, R.N.) 

Fig. 2. Rot-hole in beech tree (vide Aigburth Vale record). 

Fig. 3. Hole formed in the exposed root of a beech tree, Burnham 
Beeches, Bucks. (Photograph by Miss M. Carter.) 

Fig. 4. Closer view of fig. 3. (Photograph by Miss M. Carter.) 

Note. —In all the figures the black arrows are directed towards^ the 
orifices of the rot-holes; in fig..2/ the white line indicates 
approximately the size of the cavity. 






45 * 


Plate XII 

Larvae of the commoner English tree-hole breeding mosquitoes. 

Fig. i. Anopheles plumbeus, Steph. Larva in normal resting 
position at surface, (x 18 circa.') 

Fig. 2. Do. do. (x 6 circa.) 

Fig. 3. Anopheles plumbeus , Steph. Larva in position not 
uncommonly assumed at surface, (x 18 circa.) 

Fig. 4. Do. do. (x 6 ctrca.) 

Fig. 5. Ochlerotatus geniculatus, Ol. Larvae resting at surface. 

(x 6 circa.) 

Fig. 6. Do. Larva, (x 18 circa.) 









♦S 3 


DESCRIPTIONS OF THE MALE 
GENITAL ARMATURES OF THE 
BRITISH ANOPHELINE MOSQUITOES 

BY 

HENRY F. CARTER 
(Received for publication 16 February , 1920) 


GENUS ANOPHELES, Meig. 

Anopheles maculipennis , Meig. 

(Figs. 1 and 2) 

Basal portion of clasper more than twice as long as wide, broad 
and gently rounded distally. Dorsal surface clothed with hairs 
which are of moderate length on the outer side and at the apex. 
Ventral surface with scattered hairs on the sides and apical half, 
and two stout tuberculate spines at the base. Inner margin with a 
stout recurved bristle arising just beyond the middle. Terminal 
portion of clasper rather stout, about one and a quarter times as long 
as the basal portion, with very fine lateral hairs on the inner side, 
and a patch of very minute ones at the base. Apical tooth relatively 
short and broad. 

Basal lobes (harpagones) relatively small, each composed of two 
portions; the larger inner portion clothed with short hairs and 
bearing on its apical margin a short stout hair or spine, a shorter, 
finer hair and two short pointed spines or bristles; the smaller 
portion, a tubercle, lying above the outer side of the main area, 
carrying one or two short blunt spines. 

Penis sheath very long and narrow, with three almost straight 
leaflets on each side at its extremity; the apical leaflet is very long 
and extends as far as the tips of the spines arising from the basal 
lobes. Ninth segment with two conspicuous, slightly curved, rod¬ 
like, chitinous processes projecting backwards from the ventral 
surface. Anal membrane hairy. 








455 

Anopheles bifurcatus, L. 

(Figs. 2 and 3) 

Basal-portion of clasper about two and a half times as long as 
broad, sub-conical; apex broadly rounded. Dorsal surface with 
numerous short hairs on the inner half and many very long coarse 
hairs on the outer half and near the apex. Ventral surface with 
sparsely arranged hairs on the distal half (on the outer side they are 
of considerable length), a strong slightly recurved tuberculate.spine 
and two vertical dendriform bristles at the base (see fig. 3). Inner 



Fig. 3. Anopheles bifurcatus , L. Male genitalia (x i6o circa). 


margin with a strong bristle, recurved at the tip, arising shortly 
before the apex. Terminal portion of clasper relatively stout, rather 
longer than the basal portion and bearing beside the usual hairs a 
group of fine ones near the base. Apical tooth short and stout. 




Basal lobes large and complex, each consisting of three divisions; 
the innermost division is clothed with minute hairs and bears a large 
tubercle externally from which arise two moderately stout curved 
bristles and, between the latter, a short hair; the second portion 
lies below the third and bears two closely apposed* unequal 
chitinous processes, the tips of which are composed of much thinner 
chit in and appear recurved when mounted without pressure; third 
division with three flattened spear-shaped spines. 

Penis sheath elongate with eight short, slender terminal leaflets. 
Ninth segment without ventral processes. Anal membrane hairy. 


Anopheles ( Coelodiazesis ) pltimbeus> Stephens. 

(Figs. 2 and 4) 

Basal portion of clasper rather more than twice as long as broad, 
sub-conical with the apex broadly rounded. Dorsal surface with 
scattered hairs which are longer on the outer side and near the apex. 
Ventral surface with short scattered hairs, confined chiefly to the 



Fig. 4. AwfhtUt plumb*us , Steph. Male genitalia (X 160 circa.) 


outer side and apical half, and two strong basal spines with recurved 
tips arising from adjacent tubercles. Inner margin with a strong 
bristle, recurved at the apex, situated slightly beyond the middle. 




457 


Terminal portion of clasper moderately stopt, about one- and a 
quarter times the length of the basal portion with sever.al fine lateral 
hairs on the inner side and three on the outer side near the apex. 
Apical tooth slender. 

Basal lobes moderately large, the inner portion partly clothed 
with minute hairs and with a conspicuous hair and a long somewhat 
flattened curved bristle projecting from the apical margin; outer 
and ventral portion in the form of a small lobe or tubercle, bearing 
three or four curved, flattened, distally expanding spines or bristles. 

Penis sheath short and broad; without terminal leaflets. Ninth 
segment without ventral processes. Anal membrane covered with 
small hairs. 




459 


ON HUMAN TRYPANOSOMIASIS IN 

PERU 


BY 

WARRINGTON YORKE, M.D., 

PROFESSOR OF PARASl TOI.OC.V, SCHOOL OF TROPICAL MEDICINE, UNIVERSITY OF LIVERPOOL 


(Received for publication February 28, 1920) 

In December last, Eseomel (1919) recorded that he had discovered 
trypanosomes in the blood of a patient coming from the tropical forests 
in the eastern portion of Peril bordering on Brazil and Bolivia. As this is 
the first recorded case of human trypanosomiasis in Peru, the observation 
is not without interest. The following is a summary of the clinical 
account of the case given by Eseomel:— 

‘The patient stated that he had. for a long time, suffered from forest fevers 
of variable duration and intensity. At the time he was seen by Eseomel the 
condition was sub-febrile, the pulse was feeble, and there was general pallor with 
the greenish tint typical of the inhabitants of the region in the centre of equatorial 
America. The eyelids, the limbs, and the rest of the body exhibited a condition 
of solid oedema, which, as there was neither valvular disease of the heart nor 
albuminuria, was really a myxoedema. The patient complained of great lassitude 
and extreme prostration, and stated that for some time he had suffered from 
uncontrollable desire to sleep, which overcame him no matter where he was. 
'The reflexes were slightly decreased, and the appetite was very poor; the spleen 
and liver were hypertrophied.’ 

In view of the locality where the disease was contracted—the region 
bordering on Brazil where Triatomei megisia and other vectors of trypano¬ 
somiasis exist- -and of the symptoms exhibited, viz., myxoedema, 
previous febrile attacks followed by the sub-febrile condition, the great 
prostration and uncontrollable desire to sleep, Eseomel considered it 
necessary to examine the blood of the patient. His research was rewarded 
by the discovery of a trypanosome, which, he states, is probably 
Schizotrypanosom critzi, and adds that the existence of this trypanosome 
is not astonishing in the region where the disease was contracted. 

The following is EscomeEs account of the parasite based on examina 
tion of dehaemoglobinised thick films stained with Giemsa or Leishman :— 

‘Avec le Giemsa, ils se presentment teints en bleu clair avec le flagellc, le 
noyau et lc blepharoplaste en rouge pourpre. Le protoplasma des leucocytes cst 
bleu clair, leurs noyaux ct les hematoblastes pourpre fonce. Le fond de la 
preparation cst rose tres pale en raison des restes des hematics hjinolysees.* 



460 


Les parasites se montraient tres nets; onduleux, serpigineux, avec une longeur 
variant entre 20 et 40 fi, flagelle compris; largeur maxima, 3 a 4/<. Le flagelle 
ctait toujours plus long que le corps protoplasmique. 

Celui-ci est granuleux sur toute son etendue. La membrane ondulante a un 
aspect variable, suivant la position dans laquelle le parasite a ete fixe. 

Le noyau violet-pourpre au Giemsa, siege presque toujours a la hauteur de la 
moitie du corps. 

Le blepharoplaste, petit, a peine visible, mais toujours bien differcncie, se 
continue par le flagelle.’ 

Examination of this description reveals certain peculiarities which 
lead one to doubt whether Escomers parasite is really Schiz. cruzi; 
unfortunately, the text figure illustrating the trypanosome is not 
sufficiently good to afford any aid in forming an opinion. 

Two points in the description of the parasite call for special attention. 
In the first place, the size of the Peruvian trypanosome, viz., 20-40/4 in 
length and 3-4/4 in breadth, is very much greater than that of Schiz- 
entzi , the length of which in the blood averages about 20/4, certainly 
never approaching 40/4, and the breadth of which is not more than 2-3/4; 
and, in the second place, the ‘ small, hardly visible blepharoplast 1 in no 
way resembles this structure in Schiz. cruzi where it is always voluminous 
and assumes either an elongate or, more usually, an ovoid form. 

In these two respects, then, Escomel's trypanosome presents striking 
differences from Schiz. cruzi. Of course, it must be borne in mind that 
Escomel based his description on the examination of thick blood films, 
but this fact can hardly explain the great size of the Peruvian parasite 
and the minuteness of its blepharoplast.. 

An interesting feature in the symtomatology of the case, on winch 
considerable stress is laid by Escomel, is the overpowering somnolence 
from which the patient suffered. So far as I am aware this symptom has 
not been noted in Chagas' disease, although, of course, it is a striking 
feature of African trypanosomiasis of man. 

Assuming Escomel’s description to be correct, one is compelled 
to conclude that the trypanosome found by him in the inhabitant of the 
eastern Peruvian forests differs from Schiz. cruzi and is in reality a new 
species. As the parasite in question is capable of infecting man, tlu* 
question is important and one which requires further investigation. Tn 
the meantime, I propose for this Peruvian parasite of man the name of 
Trypanosoma escomeli in recognition of its discoverer. 

REFERENCE 

KsroMir , K. (11JI9). Hull. Sor. Path. F.xot ., Vol. XXI, p. 72 



INDEX 




INDEX 


PAGE 


Index of Authors . iii 

General Index . iv 

Index of Species and Varieties new to Science . x 


INDEX OF AUTHORS 


PAGE 

Blacklock, B. 297 

Blacklock, B.; and Carter, H. F. 187,413, 

421 

Blacklock, B. ; and O’Farrell, W. R. 189 

Blacklock, B.; and others 63, 69, 73, 75, 
97, 101, 117, 119, 125 

Breinl, A ; and Young, W. J. 351 

Carter, H. F. 453 

Carter, H. F.; and Blacklock, B. 187,413, 

421 

Carter, H. F.; and others 63, 69, 73, 75 

Carter, H. R. 299 

Cooper, C. F.; and others 63, 69, 73, 75 

Evans, A. M. 31 

Fielding, J. \V. 259 

Hatori, J. 233 

Lipkin, I. J. 149 

Macfie, J. W. S. 23, 95, 343, 347 

Macfie, J. W. S. ; and Yorke, \V. 57, 133, 

137 . 


PAGE 

Macfie, J. \V. S. ; and others 63, 69, 73, 

75 , 97 > ii 7 > 125 

Matthews, J. R. 17 

Matthews, J. R.; and Smith, A. M. 83, 91 

Morris, H. M. 339 

O’Farrell, W. R.; and Blacklock, B. 189 

O’Farrell, W. R.; and others 117,119,125 

Rodhain, J. 109 

Scott, H. H... 195 

Smith, A. M. 1, 177 

Smith, A. M.; and Matthews, J. R. 83, 91 
Stephens, J. W. W.; and others 63, 69, 73, 
75,97,101,117/119,125 

Yorke, VV. 459 

Yorke, W.; and Macfie, J. W. S. 57, 133, 

137 

Yorke, W.; and others 63, 69, 73, 75, 97, 
101,117,119,125 


Young, W. J...215, 313 

Young, W. J.; and Breinl, A. 351 


GENERAL INDEX 


PAGE 

Albuminuria in tropical Australia ... 230 

American horses, Strongylidae in . 137 

Amoeba Umax , phagocy tosis of erythro¬ 
cytes by . 133 

Amoebic dysentry in asylum patients 177 
Ancylostoma ceylanicum in the cat in 

Durban. 297 

Anopheles bifurcatus y experimental in¬ 
fection of, in 
England, with 

P. vivax . . 4*3 

„ „ male genital armature 455 

„ „ synonymy of . 422 

„ maculipennisy male genital 

armature. 453 


PAGE 

Anopheles plumbeuSy adult habits . 424 

„ „ breeding-places... 426 

„ „ as carrier of 

malaria . 438 

„ „ distribution in 

British Isles ... 435 
„ „ experimental in¬ 

fection of, with 
P. vivax ...187,413 
„ „ general distribu¬ 


tion . 423 

larval habits . 432 

male genital arma¬ 
ture . 456 

morphology . 423 


iii 


































PAGE 


PAGE 


Anopheles plumheus , observations on 
breeding-places, 
occurrence in 
Liverpool dis¬ 
trict and possible 
connection with 
spread of malaria 421 


„ „ synonymy of . 422 

Anophelines, British, male genital 

armatures of. 453 

Arsenic in treatment of malaria, 75,97,101, 

u 9 

Ascaris lumbrtcoides in convalescent 

dysenteries . 89 

Asylum patients, amoebic dysentery in 177 
„ „ intestinal infections 

in . 91 

Australia, Northern, Climate of . 353 

„ Tropical^ and its settlement 351 
Bacillus paratyphosus, coincident with 

malaria . 197 

„ typhosus , coincident with 

malaria ..’. 197 

Bionomics of Stegomyia fasciata . 259 

Blacklock, B. Ancylostoma ceylanicum 
in the cat in Durban . 297 


Blacklock, B., and Carter, H. F. The 
experimental infection, in England, 
of Anopheles plumheus , Hal., with 
Plasmodium vivax. (Sporozoites in 

salivary glands.) . 187 

Blacklock, B., and Carter, H. F. The 
experimental infection, in England, 
of Anopheles plumheus , Stephens, and 
Anopheles hifurcatus , L., with Plas¬ 
modium vivax . 413 

Blacklock, B., and Carter, H. F. 
Observations on Anopheles ( Coelo - 
diazesis) plumheus , Stephens, with 
special reference to its breeding- 
places, occurrence in the Liverpool 
district, and possible connection 


with the spread of malaria. 421 

Blacklock, B., and O’Farrell, W. R. 
Note on a case of multiple infection 

by Dracunculus medinensis . 189 

Blacklock, B., and others. Studies in 
the Treatment of Malaria, 63, 60, 73, 
75,97, 101, 117, 119, 125 

Blackwater, quinine in case of . 169 

Blood conditions of Europeans in 

tropics . 377 


Blood pressure in tropics . 377 

Brcinl, A., and Young, W. J. Tropical 

Australia and its settlement . 351 

British Anopheline mosquitoes, male 

genital armatures of. 45 3 

j Carpoglyphus anonymus . 339 

Carter, H. F. Descriptions of the 
male genital armatures of the British 
Anopheline mosquitoes . 45 3 


Carter, H. F., and Blacklock, K. The 
experimental infection, in England, 
of Anopheles plumheus , Hal., with 
Plasmodium vivax. (Sporozoites in 

salivary glands.) . 187 

Carter, H. F., and Blacklock, B. The 
experimental infection, in England, 
of Anopheles plumheus , Stephens, and 
Anopheles hifurcatus , L., with Plas¬ 
modium vivax . 413 

Carter, H. F., and Blacklock, B. 
Observations on Anopheles ( Coelo - 
diazesis) plumheus , Stephens, with 
special reference to its breeding- 
places, occurrence in the Liverpool 
district, and possible connection 

with the spread of malaria. 421 

Carter, H. F., and others. Studies in 
the Treatment of Malaria, 63, 69, 73, 75 
Carter, H. R. The mechanism of the 
spontaneous elimination of yellow 

fever From endemic centres . 299 

Cat, Ancylostoma ceylanicum in the ... 297 
Cephalophus sylvicultor , Filaria per - 

ttnue in. 109 

Chilomastix mesnili in convalescent 

dysenteries, 88, 92, 

i8 4 

„ „ population of 

Great Britain, 92, 
184 


Climate of Northern Australia . 353 

Clothing in the tropics ..*... 391 

Coincident malaria and enteric fever 195 
Composition of the urine in the 

tropics . 21 q 

Contribution to the question of the 
number of races in the species 

Entamoeba histolytica . I 

Cooper, C. F., and others. Studies in 
the Treatment of Malaria, 63, 69, 73, 75 
Crescents, Disappearance of, under 
quinine treatment . 73 


IV 


































PAGE 

Cylicostomum pateratum , sp.n. 57 

„ spp. in horses .138,139 

„ „ pathogenicity of ... 142 

Dermofilariose cheloidiforme chez 

Cepbalophus sylvicultor . 109 

Dironlaria kuelzii . 113 I 

Distribution and destruction of quinine ] 

in animal tissues . 149 

Draeunculus medinensis , Case of mul¬ 
tiple infection by . 189 

Durban, Ancylostoma ceylanicum in the 

cat in . 297 

Dysenteries, Convalescent, Intestinal 

infections among. 83 

Dysentery, Amoebic, in asylum 

patients . 177 

Emetine, alleged effect on size of E. 

histolytica cysts. 6 

Endemic Tsutsugamushi disease of 

Formosa ... 233 

Entamoeba coli in convalescent dysen¬ 
teries .88, 92, 184 

„ „ The course and dura¬ 

tion of an infection 

with . 17 

„ „ in population of Great 

Britain .92, 184 j 

,, histolytica , Alleged effect 
of emetine on 
size of cysts... 6 

„ „ in convalescent 

dysenteries 87, 92, 
184 

,, „ measurements of 

cysts. 2 

,, ,, Number of races 

in the species 1 

,, „ in population of 

Great Britain 92,184 
nana in convalescent dysen¬ 
teries .84, 92, 184 

„ „ in population of Great 

Britain .92, 184 

Enteric fever, coincident with malaria 195 
Evans, A. M. On the genital arma¬ 
ture of the female tsetse-flies 

( Glossina) . 31 

Experimental infection, in England, of 
Anopheles bifurcatus with P. vivax ... 413 
„ plumbeus with P . vivax 187,413 j 

Fielding, J. W. Notes on the biono¬ 
mics of Stegomyia fas data . 259 


PACE 

Filaria perienuty n.sp., provoquant une 
dermofilariose cheloidiforme chez 


Cephalopbus sylvicultor . 109 

Formosa, Endemic Tsutsugamushi 

disease in . 233 

Genital armature of female Glossina 31 


„ „ „ male Anophelinae 45 3 

Giardia intestinalis in convalescent 

dysenteries 88, 92, 184 
„ „ in population of 

Great Britain 92, 184 


Glossina , genital armature of female 31 

„ austenty female armature. 55 

„ brevipalpisy female armature 48 

„ caliginea , female armature 52 

„ fusca y female armature . 37 

„ „ systematic table of 

group. 36 

„ fuscipleurisy female armature 40 

„ longipennisy female armature 40 

„ medicorumy female armature 46 

„ morsitansy female armature... 55 

„ nigrofusedy female armature 39 

„ pallicerdy female armature ... 52 

„ palpalisy female armature ... 50 

„ s ever ini y female armature ... 44 

„ tabaniformisy female armature 42 

„ tachinoidesy female armature 53 

Gold Coast, Xerophthalmia in native 


of . 343 

GyalocephaluSy spp. in horses .138, 139 

Hatori, J. On the endemic Tsutsu¬ 
gamushi disease of Formosa . 233 

Heat, Effects of, on animal organism... 368 

Housing in tropical Australia . 403 

Human trypanosomiasis in Peru . 459 

Humidity, Effects of, on animal 

organism . 368 

Hypopus of Carpoglyphus anonymus ... 339 
Influence of external temperature and 
rate of cooling upon the respiratory 

metabolism . 313 

Intestinal infections among convales¬ 
cent dysenteries. 83 

„ protozoal infections in 

Asylum patients . 91 

„ protozoal infections in Uni¬ 
versity and school cadets 94 
Iodine cysts in convalescent dysenteries 89 
Katathermometer, for measuring rate 

of heat loss .*. 320 

Leukaemia, associated with malaria ... 347 


v 




































PACK 


Lipkin. I. J. On the distribution and 
destruction of quinine in animal 

tissues . 149 

Liquor arscnicalis in treatment of 

malaria . 119 

Macfie, J. W. S. An observation on 
the effect of malaria in leukaemia ... 347 
Macfie, J. W. S. Presentation of the 

Mary Kingsley Medal to . 95 

Macfie, J. VV. S. Two parasites of 

Naja nigricollis . 23 

Macfie, J. \V. S. Xerophthalmia in a 

native of the Gold Coast . 343 

Macfie, J. W. S., and Yorke, W. The * 
phagocytosis of erythrocytes by an 
amoeba of the Lmax type . 133 


Macfie, J. VV. S., and Yorke, W. 

Strongylidae in Horses — 

VII : Cylirostomum pateratum , sp. n. 57 
VIII: Species found in American 

horses. 137 

Macfie, J. \V r . S., and others. Studies 
in the Treatment of Malaria, 63, 69, 73, 
75,97, 101, 117, 119, 125 

Malaria, A. plumbeus as carrier of. 421 

,, coincident with enteric fever 195 

,, in leukaemia. 347 

,, Studies in the Treatment of, 63, 
69, 73 . 75 . 97 . *°i» 11 7 . 

119. I2 5 

Malignant tertain malaria, Treatment 

of ..63,69,73,75 

Mary Kingsley Medal, Presentation of, 

to Dr. J. \V. S. Macfie . 95 

Matthews, J. R. The course and 
duration of an infection with 

Entamoeba coli . 17 

Matthews, J. R., and Smith, A. M. 

The intestinal protozoal infections 
among convalescent dysenteries 
examined at the Liverpool School 
of Tropical Medicine. (Third 

Report.) . 83 

Matthews, J. R., and Smith, A. M. 

The spread and incidence of intes¬ 
tinal protozoal infections in the 
population of Great Britain. IV : 
Asylum patients. V: University 


and school cadets . 91 

Mechanism of the spontaneous elimina¬ 
tion of yellow fever from endemic 

centres . 299 


PAC.F 

Metabolism of white races living in 
the tropics :— 

II : The composition of the urine ... 215 

III : The influence of external tem¬ 

perature and rate of cooling 
upon the respiratory metabo¬ 


lism . 313 

Morris, H. M. The hypopus of 
Carpoglyphus anonymm , Haller .. •••• 339 
Mosquitoes, methods of keeping and 

feeding .:. 260 

Naja nigricollis , Two parasites of. 23 

Nervous system, Effect of tropical 

climate on . 389 

Nitrogen in urine . 223 

Novarsenobillon in treatment of 

malaria.99, 101 

Observation on the effect of malaria in 

leukaemia . 347 

Oesopbagodontus robustus in horses ... 139 
O’Farrell, W. R., and Blacklock, B. 

Note on a case of multiple infection 

by Dracunculus medinensis . 189 

O’Farrell, W. R., and others. Studies 


in the Treatment of Malaria, 117, 119, 

125 

Oxyuris vermicularis in convalescent 


dysenteries . 89 

Peru, Human trypanosomiasis in . 459 

Phagocytosis of erythrocytes by an 

amoeba of the Limax type... 133 

Phosphates in urine . 222 

Physiological changes of white man in 

tropics.372, 394 

Plasmodium mesnili , in a snake . 25 


,, vivax infecting A. bifur- 

catus ... 413 
„ „ 4 . plum- 

beus 187, 413 

Protozoal infections among convales¬ 
cent dysenteries 83 

„ „ in the population 

of Great Britain 91 

Quartan malaria, Treatment of. 97 

Quinine, Action of liver on . 161 

„ in case of blackwater fever... 169 

„ distribution and destruction 

of, in animal tissues . 149 

„ bihydrochloride in treatment 

of malaria, 63, 97, 101,119, 125 

„ sulphate in treatment of 

malaria .69,73,125 


vi 





































PAGE 


(Juitenine hydrochloride in treatment 

of malaria 117 

„ „ urine. 169 

Rumsden, W. Preliminary report on 
liquids from cavities of hollow 

trees . 440 

Relapses in simple tertian malaria. 125 

Respiration, Rate of, in tropics. 376 

Respiratory metabolism, influence of 
external temperature and rate of 

cooling . 313 

Rodhain, J. Filaria pertenue , n. sp., 
provoquant une dermofilariose 
cheloldiforme chez Cephalophus syl- 

vicultor . 109 

Scott, H. H. Coincident malaria and 
enteric fever . 195 


Simple tertian malaria, Treatment of, 101, 

117,119,125 

Smith, A. M. Cases of acute amoebic 
dysentery in asylum patients never 

out of England . 177 

Smith, A. M. A contribution to the 
question of the number of races in 
the species Entamoeba histolytica ... I 
Smith, A. M., and Matthews, J. R. 

The intestinal protozoal infections 
among convalescent dysenteries 
exatnined at the Liverpool School 
df Tropical Medicine. (Third 

Report.) :. 83 

Smith, A. M., and Matthews, J. R. 

The spread and incidence of intes¬ 
tinal protozoal infections in the 
population of Great Britain. IV : 
Asylum patients. V: University 

and school cadets. 91 

Spread and incidence of intestinal 
protozoal infections in the popula¬ 
tion of Great Britain. IV : Asylum 
patients. V : University and school 


cadets . 91 

Sugomyia , extermination unnecessary 
for elimination of yellow 

fever . 299 

„ fasciata , adults . 292 

„ „ Bionomics of . 259 

„ „ effect of light on 

larval develop¬ 
ment . 282 

,, ,, eggs. 261 

,, „ larvae . 282 


PAGE 

Stegomyia Jasa'dtd, quantity and nature 
of food influencing 
larval development 285 
Stephens, J. W. YV., and others. 
Studies in the Treatment of Malaria, 63, 
69, 73. 75, 97. ioi, 117, 119, 125 

Strongylidae in Horses.57, 137 

Strongyloides in convalescent dysenteric 89 

Strongylus spp. in horses .138, 139 

„ „ Pathogenicity of . 142 

Studies in the Treatment of Malaria— 
XXII. Intramuscular injections of 
quinine bihydrochloride 
grains 15 on each of two 
consecutive days only, in 
malignant tertian malaria 63 
XXII 1 . Oral administration of 
quinine sulphate grains 30 
on each of two consecutive 
days weekly, over a period 
of five weeks, in malignant 

tertian malaria . 69 

XXIV. The disappearance of cres¬ 
cents under quinine treat¬ 
ment . 73 

XXV. Arsenic in malignant tertian 

malaria . 75 

XXVI. The action of arsenic and of 
quinine on quartan 

malaria . 97 

XXVII. Intravenous injections of 
novarsenobillon and intra¬ 
muscular injections of 
quinine bihydrochloride 
in simple tertian malaria 101 
XXV III. Quitenine hydrochloride in 

simple tertian malaria ... 117 
XXIX. Oral administration of 
liquor arsenicalis minims - 
30 daily for sixteen days, 
with quinine bihydro¬ 
chloride grains 15 intra¬ 
muscularly on the first and 
second, eighth and ninth, 
fifteenth and sixteenth 
days, in simple tertian 

malaria . 119 

XXX. At what time after cessation 
of quinine treatment do 
relapses occur in simple 
tertian malaria ? (Second 
communication.) . 125 





























PACK 


PAGE 


Sunlight in the Tropics . 363 

Sweat, Amount of water and sodium 

chloride lost in . 219 

Taenia in convalescent dysenteries ... 89 

Tartar emetic in leukaemia . 347 

Temperature of body in tropics, 369, 374 
„ external, influence on 

respiratory metabolism 313 

„ 4 skin-shirt 9 . 318 

Treatment of malaria, 63, 69, 73, 75, 97, 
101,117,119,125 

Trichomonas intestinalis in convales¬ 
cent dysenteries . 84 

Trichuris trichiura in convalescent 

dysenteries . 89 

Triodontopkorus spp. in horses.138, 139 

Trombicnla akamushi . 241 

Tropical Australia and its Settlement 351 

Tropics, Blood conditions of Europeans 

^ . 377 

„ Blood pressure in . 377 

„ Clothing in . 391 

„ Metabolism of white races 

living in .215, 313, 380 

„ Physiological changes of white 

man in . 394 

,, Rate of respiration in . 376 

Sunlight in . 363 

Trypanosoma escomeli , sp.n. 459 

„ najae . 24 

„ primeti . 24 


„ voltariae , sp.n. 23 i 

Trypanosomiasis, human, in Peru ... 459 
Tsetse-flies, Genital armature of female 31 
Tsutsugamushi disease of Formosa ... 233 
„ „ Causal 

organism of 240 

„ „ diagnosis . 249 

„ „ prophylaxis ... 252 ; 


Tsutsugamushi disease, symptomat¬ 
ology . 246 

,, „ treatment ... 249 

University and school cadets, Intestinal 

infections in. 94 

Urine, Composition of, in the tropics 215 

„ Inorganic salts in . 219 

„ Nitrogen in .. 223 

„ Phosphates in .. 222 

White races living in the tropics, 


Metabolism of ...215, 313, 380 


„ settlement in tropical Australia 397 
Xerophthalmia in a native of the Gold 

Coast . 343 

Yellow fever, Spontaneous elimination 

of . 299 

Yorke, W. On human trypanosomiasis 

in Peru. 459 

Yorke, W., and Macfie, J. W. S. The 
phagocytosis of erythrocytes by an 

amoeba of the Limax type . 133 

Yorke, W., and Macfie, J. W. S. 
Strongylidae in Horses VII: Cyli- 
costomum pater atum , 

sp-n. 57 


VIII: Species found 
in American horses 137 
Yorke, \V., and others. Studies in the 
'Treatment of Malaria 63, 69, 73, 75, 
97, 101, U7, 119, 125 
Young, W. J. The metabolism of 
white races living in the tropics:— 

II. The composition of the urine ... 215 
III. The influence of external tem¬ 
perature and rate of cooling 
upon the respiratory meta¬ 


bolism . 313 

Young, W. J., and Breinl, A. Tropical 
Australia and its Settlement . 351 


INDEX OF SPECIES NEW TQ SCIENCE 

PACE j PAGE 

Cylicostomum pater atum ... ... 57 Trypanosoma escomeli . ... 459 

Filaria pertenue ... ... ... 109 j Trypanosoma voltariae ... ... 23 


VUl